GIFT  OF 


Charles 


EARTH 

SCIENCES 

LIBRARY 


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MORPHOLOGY    AND    TELEOLOGY, 


ESPECIALLY     IN     Til  1C 


LIMBS    OF    MAMMALIA. 


13  Y  HURT  G.  WILDER,  S.B. 


I  KItOM    TilK    MKJIOlJtS    Kl'.Al)    1;]<:H>KK    TIIK    IIOWTO.N    SOCIETY    OI     NATl!14AL    HISTOKY,    VOL.    I.    SO.    I.) 


CAMBRIDGE: 

PRINTED  AT   THE    RIVERSIDE   PRESS. 

NOVEMBER,    180;"). 


ON 

MORPHOLOGY   AND    TELEOLOGY, 


ESPECIALLY     IN     THE 


BY  BURT  G.  WILDER,  S.B. 


|  FROM    THK   MKMOIRS    HEAD    BEFOKE  THE   BOSTON    SOCIETY   OF   NATURAL   HISTORY,    VOL,.    I.   NO.   I.] 


CAMBRIDGE: 
PRINTED  AT   THE   RIVERSIDE   PRESS. 

NOVEMBER,   1865. 


SCIENCE 
LIBRAW 


ON  MORPHOLOGY  AND  TELEOLOGY,  ESPECIALLY  IN  THE  LIMBS  OF  MAMMALIA. 

BY  BURT   G.   WILDER,   S.  B. 

Read  June  3d,  1863.  /  '-     , 


IT  is  not  many  years  since  the  very  title  of  this  paper  would  have  been  enough  to  'insure 
its  remaining  unread  by  most  professional  men,  or,  if  read,  to  excite  their  derision  of  him 
who  should  have  so  wasted  his  time  as  to  write,  or  even  think,  of  such  vain  abstractions,  fit 
expressions  of  the  useless  imaginings  of  the  half-crazy  enthusiast  Oken,  and  his  only  less 
crazy,  because  less  gifted,  disciples.  And  there  are,  even  now,  stern  votaries  of  practical 
science  who  would  scorn  any  attempt  to  raise  their  eyes  above  the  mere  facts  of  Nature 
which  are  as  patent  to  the  ignorant  vision  as  to  their  own,  and  who  refuse  to  seek  an 
insight  into  those  hidden  relations,  for  the  correct  understanding  of  which  their  superior 
knowledge  might  be  the  surest  preparation. 

But  there  are  others,  and  their  number  is  increasing,  who,  believing  in  the  existence  of  a 
general  plan  underlying  all  the  more  external  phenomena  of  Nature,  are  willing  to  try  to 
comprehend  it  in  its  greater  and  lesser  manifestations ;  and  they,  in  reading  the  "  Physio- 
philosophy,"  may  .be  able  to  discern,  amongst  much  that  is  fanciful  and  absurd,  many  sug- 
gestions of  a  sound  as  well  as  original  and  striking  philosophy.  No  apology,  therefore,  is 
now  required  for  thinking  or  writing  upon  subjects  which  have  engaged  the  attention  of 
the  most  celebrated  students  of  both  animal  and  vegetable  anatomy,  and  which,  I  am  con- 
vinced, will,  erelong,  be  acknowledged  to  be  as  essential  to  the  proper  understanding  of 
these  sciences  as  the  classifications  of  which  they  form  the  only  true  basis. 

To  express  the  various  relations  which  have  been  observed  among  the  several  parts  and 
their  functions,  of  animals  and  plants,  the  following  terms  have  gradually  come  into  use  : 
homology,  affinity,  morphology,  analogy,  teleology ;  to  these  may  be  added  physiology,  which, 
though  a  term  long  employed  to  denote  the  general  study  of  function,  has  now  acquired 
a  certain  technical  significance,  equivalent  to  the  more  strictly  scientific,  and  therefore 
preferable  term,  teleology. 

Analogy  is  used  to  indicate  similarity  of  function,  which  may  be  very  close,  when  yet  the 
two  parts  are  widely  dissimilar  in  structure ;  as,  for  instance,  the  organs  of  aerial  locomotion 
of  a  bird  and  a  butterfly,  which  both  go  by  the  name  of  wings,  though  one  is  built  upon  the 
vertebrate,  and  the  other  upon  the  articulate  plan  of  structure.  Of  course  the  structure 
may  correspond  with  the  external  form  and  function,  and  then  the  analogy  is  more  com- 
plete, as  between  the  foot  of  man  and  that  of  a  bear. 

Now  the  general  function  or  use  of  a  part  is  its  physiology  ;  the  special  or  principal  use  of 
a  part  is  its  final  cause  or  end,  or  teleology ;  and  parts  which  are  teleologically  similar  are 
said  to  be  analogous. 

It  is  evident  that  the  external  form  and  the  function  must  to  a  great  extent  correspond,  at 
least  much  more  fully  than  either  may  with  the  internal  structure,  and  here  we  observe  the 
first  distinction  between  the  two  groups  of  terms  given  above ;  for  this  intimate  structure 
and  arrangement,  in  other  words,  the  pure  anatomy  of  anything,  is  its  morphology,  and  parts 
which  are  morphologically  similar  are  said  to  be  homologous  ;  there  is  homology  or  affinity,  or, 
in  still  plainer  words,  more  or  less  identity  of  structure  between  them ;  and  here  again,  as  was 
seen  in  speaking  of  analogy,  parts  or  organs  which  are  homologous,  that  is,  identical  in  their 

1 


89O41 6 


ON  MORPHOLOGY  AND  TELEOLOGY,  ESPECIALLY  IN  THE  LIMBS  OF  MAMMALIA. 

BY  BURT   G.   WILDER,   S.  B. 

Kead  June  3d,  1863.  .'"'     . 

T  '*'•''>:..  •  ;•'•";;• 

J.T  is  not  many  years  since  the  very  title  of  this  paper  would  have  been  enough  to 'insure 

its  remaining  unread  by  most  professional  men,  or,  if  read,  to  excite  their  derision  of  him 
who  should  have  so  wasted  his  time  as  to  write,  or  even  think,  of  such  vain  abstractions,  fit 
expressions  of  the  useless  imaginings  of  the  half-crazy  enthusiast  Oken,  and  his  only  less 
crazy,  because  less  gifted,  disciples.  And  there  are,  even  now,  stern  votaries  of  practical 
science  who  would  scorn  any  attempt  to  raise  their  eyes  above  the  mere  facts  of  Nature 
which  are  as  patent  to  the  ignorant  vision  as  to  their  own,  and  who  refuse  to  seek  an 
insight  into  those  hidden  relations,  for  the  correct  understanding  of  which  their  superior 
knowledge  might  be  the  surest  preparation. 

But  there  are  others,  and  their  number  is  increasing,  who,  believing  in  the  existence  of  a 
general  plan  underlying  all  the  more  external  phenomena  of  Nature,  are  willing  to  try  to 
comprehend  it  in  its  greater  and  lesser  manifestations ;  and  they,  in  reading  the  "  Physio- 
philosophy,"  may-be  able  to  discern,  amongst  much  that  is  fanciful  and  absurd,  many  sug- 
gestions of  a  sound  as  well  as  original  and  striking  philosophy.  No  apology,  therefore,  is 
now  required  for  thinking  or  writing  upon  subjects  which  have  engaged  the  attention  of 
the  most  celebrated  students  of  both  animal  and  vegetable  anatomy,  and  which,  I  am  con- 
vinced, will,  erelong,  be  acknowledged  to  be  as  essential  to  the  proper  understanding  of 
these  sciences  as  the  classifications  of  which  they  form  the  only  true  basis. 

To  express  the  various  relations  which  have  been  observed  among  the  several  parts  and 
their  functions,  of  animals  and  plants,  the  following  terms  have  gradually  come  into  use : 
homology,  affinity,  morphology,  analogy,  teleology ;  to  these  may  be  added  physiology,  which, 
though  a  term  long  employed  to  denote  the  general  study  of  function,  has  now  acquired 
a  certain  technical  significance,  equivalent  to  the  more  strictly  scientific,  and  therefore 
preferable  term,  teleology. 

Analogy  is  used  to  indicate  similarity  of  function,  which  may  be  very  close,  when  yet  the 
two  parts  are  widely  dissimilar  in  structure ;  as,  for  instance,  the  organs  of  aerial  locomotion 
of  a  bird  and  a  butterfly,  which  both  go  by  the  name  of  ivings,  though  one  is  built  upon  the 
vertebrate,  and  the  other  upon  the  articulate  plan  of  structure.  Of  course  the  structure 
may  correspond  with  the  external  form  and  function,  and  then  the  analogy  is  more  com- 
plete, as  between  the  foot  of  man  and  that  of  a  bear. 

Now  the  general  function  or  use  of  a  part  is  its  physiology  ;  the  special  or  principal  use  of 
a  part  is  its  final  cause  or  end,  or  teleology •  and  parts  which  are  teleologically  similar  are 
said  to  be  analogous. 

It  is  evident  that  the  external  form  and  the  function  must  to  a  great  extent  correspond,  at 
least  much  more  fully  than  either  may  with  the  internal  structure,  and  here  we  observe  the 
first  distinction  between  the  two  groups  of  terms  given  above ;  for  this  intimate  structure 
and  arrangement,  in  other  words,  the  pure  anatomy  of  anything,  is  its  morphology,  and  parts 
which  are  morphologically  similar  are  said  to  be  homologous  ;  there  is  homology  or  affinity,  or, 
in  still  plainer  words,  more  or  less  identity  of  structure  between  them ;  and  here  again,  as  was 
seen  in  speaking  of  analogy,  parts  or  organs  which  are  homologous,  that  is,  identical  in  their 

1 


890416 


2  WILDER  ON  MORPHOLOGY  AND  TELEOLOGY 

general  plan  of  structure,  may  be  intended  to  perform  functions  most  diverse,  and  their  out- 
ward forms  be  in  like  degree  modified.  For  instance,  the  fin-like  flipper  of  the  seal  bears 
little  resemblance  to  the  anterior  extremity  of  the  ape,  and  yet  they  are  identical  in  their 
general. structure,  —  they  are  homologous. 

.'it  trfa^'have  been  inferred,  from  what  has  been  said  above,  that  we  have  necessarily  two 
.«^st^s 'of nomenclature,  according  as  morphology  or  teleology  is  taken  as  the  basis.  For 
'"'it is  tne" latter  which  confers  common  and  popular  names  on  objects  of  Natural  History,  and 
arranges  them  in  a  way  which,  though  convenient  enough  under  ordinary  circumstances, 
utterly  fails  of  precision  for  all  scientific  purposes ;  and  the  anatomist  and  zoologist  soon 
learn  that  morphology  alone  must  be  their  guide  in  scientific  nomenclature.  Thus  the 
name  fish  is  applied  to  several  animals  in  structure  very  unlike  the  true  Pisces,  merely  be- 
cause, like  that  group  of  Vertebrates,  they  live  in  the  water :  to  certain  Radiates,  as  the 
star-fish  and  sun-fish;  to  Articulates,  as  the  cray-fish;  and,  formerly,  even  to  the  whale,  an 
air-breathing  Mammalian  Vertebrate.  So  among  Articulates,  the  monosyllable  fly  forms  the 
ending  of  the  common  names  of  many  insects,  as  butter-fly,  dragon-fly,  harvest-fly,  ichneu- 
mon-fly, members  respectively  of  the  sub-orders  Lepidoptera,  Neuroptera,  Hemiptera,  and 
Hymenoptera,  though  it  is  only  to  the  Diptera  that  the  name  fly  properly  belongs. 

These  are  zoological  ambiguities ;  anatomical  ones  are  even  more  frequent.  All  organs 
of  aerial  locomotion  are  commonly  called  wings,  whether  they  are  articulate  or  vertebrate 
in  type,  or  whether,  within  the  latter  group,  they  are  avian  or  mammalian,  as  those  of  the 
bats;  and  the  same  is  the  case  with  other  parts  and  organs,  thorax,  abdomen,  heart,  liver, 
and  stomach. 

I  could  not  well  pass  over  this  most  important  branch  of  the  subject ;  but  the  great  ne- 
cessity to  the  philosophical  naturalist  for  a  revised  anatomical  nomenclature  has  already 
been  strongly  urged  by  Professor  Agassiz,  before  the  Boston  Society  of  Natural  History.1 
Premising  that  the  members  of  the  four  great  types  have  nothing  in  common  beyond 
their  all  being  animals,  and  that,  therefore,  no  parts,  however  similar  in  function,  can  pos- 
sibly be  homologous  in  animals  belonging  to  any  two  of  these  types,  he  showed  the  propri- 
ety of  restricting  the  common  names  mouth,  stomach,  heart,  and  the  like,  to  one  of  these 
groups,  the  Vertebrates  perhaps,  and  of  applying  other  names  to  the  analogous  parts  in  the 
other  three  types.  Perhaps  the  change  should  be  even  greater  than  this ;  for,  since  these 
new  names  would  of  course  be  classical  in  their  derivation,  and  the  common  ones,  though 
scientifically  restricted,  would  in  general  discourse  retain  the  same  loose  application,  it 
would  seem  better  to  employ  new  terms  for  the  various  parts  and  organs  in  each  of  the 
four  types,  leaving  the  common  ones  as  they  are  now.  It  is  evident  that  much  remains  to 
be  done  in  this  matter  of  anatomical  nomenclature,  and  that  it  is  of  as  much  importance  to 
the  anatomist  as  are  the  names  of  the  animals  themselves  to  the  zoologist. 

Popular  descriptive  zoology  concerns  rather  the  teleological  characters  of  animals,  while 
the  strictly  scientific  and  systematic  arrangements  are  based  upon  anatomy,  and  thus  upon 
morphology. 

We  have  noticed  one  distinction  between  the  terms  given  above :  that  morphology  and 
homology  both  refer  to  structure,  while  teleology  and  analogy  both  refer  to  function.  Affin- 
ity is  merely  a  common  synonym  for  homology,  and  may  therefore  be  omitted.  And  now 
the  four  principal  terms  pair  off  on  another  basis ;  morphology  and  teleology  are  absolute 
terms,  as  it  were,  and  may  refer  to  the  structure  or  the  function  of  but  a  single  part  or  organ ; 

1  See  also  his  section  on  Morphology  and  Nomenclature,  in  Contributions  to  the  Natural  History  of  the  United  States, 
vol.  iii.  chapter  ii.  section  iv. ;  also  section  iii.  p.  69. 


IN  THE  LIMBS   OF  MAMMALIA.  3 

while  homology  and  analogy  are  the  corresponding  relative  terms,  and  necessarily  refer  to 
two  or  more  parts  or  organs  which  are  morphologically  or  ideologically  similar. 

Morphology  is  not  exactly  synonymous  with  anatomy,  for  the  latter  term  embraces 
all  the  characters  of  a  part,  external  as  well  as  internal ;  so  that,  strictly  speaking,  parts 
which  are  anatomically  similar,  are  likewise  physiologically  so.  But  morphology  refers 
rather  to  the  general  plan  of  structure  of  a  part,  without  altering  which,  great  modifica- 
tions may  be  wrought  in  its  outward  aspect,  with  reference  to  the  various  functions  it  is  to 
perform. 

In  like  manner  teleology  is  not  exactly  synonymous  with  physiology,  for  the  latter 
term  embraces  all  the  functions  which  can  be  performed  by  the  part,  the  less  as  well  as  the 
more  essential,  otherwise  the  converse  of  the  previous  proposition  would  be  true,  and  parts 
which  were  teleologically  similar  would  be  also  morphologically  similar,  which  is  not  the 
case  ;  every  form  or  morph  has  a  certain  general  use  or  function  proper  to  it,  and  which 
may  remain  under  many  of  its  modifications. 

It  is  thus  of  the  utmost  importance  to  discriminate  between  essential  structure  or 
morphology,  and  general  structure  or  anatomy ;  so  also  between  special  function  or  tele- 
ology, and  general  function  or  physiology. 

Most  objects,  whether  animal  or  vegetable,  and  their  various  organs,  possess  more  than 
one  attribute  ;  their  anatomy  is  compound ;  their  morphology  is  that  simple  essential  struc- 
ture which,  as  a  foundation,  underlies  the  more  external  attributes,  one  of  which  is  specially 
developed  for  the  performance  of  the  function  from  which  it  has  its  name ;  by  an  easy 
transfer,  the  name  is  finally  associated  in  our  mind  with  the  morphology ;  and  then,  if  this 
primary  attribute  be  overshadowed  by  an  excessive  development  of  one  of  the  secondary 
attributes,  although  the  function  of  the  part  may  be  entirely  changed,  yet,  as  the  essential 
structure  is  still  recognizable  through  the  external  mask,  the  name  is  unchanged.  Morpho- 
logically it  is  the  same,  though  teleologically  it  may  be  quite  another  thing.  For  examples, 
and,  if  I  remember  rightly,  a  clearer  explanation  of  this  transfer  of  the  name  of  a  part,  see 
the  opening  paragraphs  in  Owen's  Report  on  the  "  Homologies  of  the  Vertebrate  Skeleton," 
to  the  British  Association  for  the  Advancement  of  Science  for  the  year  1846. 

With  things  inanimate  the  teleology  is  the  use  which  is  made  of  them.  As  a  familiar 
illustration,  the  round  Dutch  cheeses,  used  as  missiles  by  one  of  the  parties  in  a  sea-fight 
years  ago,  were  none  the  less  cheeses,  and  perhaps  excellent  ones,  because  on  this  occasion 
put  in  the  place  of  round  shot,  thus  making  the  use  for  which  they  were  intended  and 
named  subservient  to  one  rendered  possible  by  a  secondary  attribute,  their  extreme  hard- 
ness. As  a  second  example,  far  too  familiar  in  these  days,  a  shell  may  strike  a  victim 
before  it  explodes,  and  thus  be  teleologically  a  solid  shot,  while  yet  its  structure,  as 
adapted  to  its  intended  use,  remains  unaltered. 

It  is  needless  to  multiply  illustrations.  Whenever  anything,  without  alteration  in  its 
essential  structure,  even  though  its  external  form  be  somewhat  modified,  fulfils  a  function 
other  than  that  for  which  it  was  originally  intended,  then  its  morphology  and  its  teleology, 
previously  coinciding,  are  at  variance. 

It  thus  appears  that  the  teleology  may  differ  from  the  morphology,  as  the  spirit  of  the 
law  from  the  letter  thereof,  as  the  expression  of  a  face  from  the  features  composing  it,  as  the 
practical  from  the  technical  or  theoretical,  as  the  actual  or  virtual  from  the  nominal  or  ostensible  ; 
in  short,  as  the  thing  may  differ  from  its  name,  the  de  facto  from  the  dejure. 

Morphology  is  substantive ;  teleology  is  adjective.  Morphology  is  the  noun ;  teleology  is  its 
modifier.  And  as  the  noun  with  its  modifier  may  be  regarded  as  a  compound  substantive, 


4  WILDER  ON  MORPHOLOGY  AND  TELEOLOGY 

and  may  thus  be  further  modified  by  other  adjectives,  so  in  comparative  anatomy,  nothing  is 
absolutely  morphological  or  teleological,  but  only  with  reference  to  some  organ  or  function 
more  general  above,  or  more  special  below ;  it  is  the  possibility  of  the  configuration  of  an 
organ  being  modified  without  change  to  a  corresponding  degree  in  the  internal  structure 
and  arrangement  of  parts.  Zoologically  speaking,  it  is  the  possibility  of  specific  modifica- 
tions of  generic  ideas,  so  that  from  a  limited  number  of  substantives,  by  adjective  additions, 
are  made  designations  of  many  more  objects ;  and  few  at  this  day  dare  affirm  that  this  is 
only  a  matter  of  human  invention  for  human  convenience. 

Every  genus  represents  the  morphology  of  the  species  embraced  within  it,  and  they 
are  teleological  modifications  of  the  generic  idea ;  now  this  is  the  relation  between  each 
higher  group  and  the  next  lower ;  the  further  we  recede  from  the  species,  from  the  indi- 
vidual in  fact,  the  more  occult  and  ideal  becomes  the  morphology,  till  we  reach  at  last  the 
four  great  groups  called  types,  which,  as  we  shall  see  hereafter,  may  even  be  represented  by 
geometrical  figures.  How  far  are  these  removed  from  the  living  sentient  individuals  which 
form  the  other  zoological  extreme  !  And  yet  it  does  not  follow  that  the  existence  of  the 
order,  the  class,  or  even  the  type,  is  any  less  real  and  actual  than  that  of  the  species  or  of 
the  individual ;  it  is  less  material.,  but  none  the  less  substantial ;  in  fact,  the  higher  the  group, 
the  more  real  and  enduring  it  is,  for  it  exists  in  all  the  members  of  all  the  groups  embraced 
within  it,  though  it  would  exist  if  it  had  but  a  single  individual  representative. 

It  was  said  above,  that  morphology  refers  only  to  the  general  plan  of  structure ;  in  a 
certain  sense  this  is  so,  since  it  refers  to  a  more  hidden  interior  grade  of  anatomical  charac- 
ters than  those  which  ordinarily  appear  upon  the  surface.  The  zoologist  will  see  that  each 
of  his  categories  of  structure  is  based  upon  a  different  grade  of  morphology ;  thus  there  is 
a  type  morphology,  the  most  interior  of  all,  beyond  which  there  are  no  homologies,  but 
within  which  are  more  and  more  apparent  ones,  the  class  homologies,  the  ordinal  homol- 
ogies, the  homologies  of  the  family,  genus,  and  species.  I  do  not  mean  to  say  that  these 
groups,  as  at  present  characterized  by  Professor  Agassiz,  or  by  any  other  naturalist,  are  the 
true  ones,  or  that  they  should  bear  these  names,  or  even  that  there  is  just  this  number  of 
categories  of  structure ;  but  I  do  believe  that  a  classification  does  exist  in  Nature  entirely 
independent  of  human  thought ;  that  the  various  kinds  of  groups  in  this  natural  classifica- 
tion are  founded  upon  categories  of  structure  radically  distinct,  not  at  all  merging  or  inter- 
changing ;  and  finally,  that  these  categories  are  simply  statements  of  the  various  grades  of 
morphology,  upon  which  alone  classifications  are  based. 

But  though  this  seems  to  carry  us  away  from  direct  material  function  or  use,  it  by  no 
means  negatives  the  idea  that  each  natural  group  does  really  represent  some  use  in  the 
grand  operations  of  Nature.  Indeed,  this  would  follow  as  the  converse  of  what  was  said 
above,  that  every  higher  group  represents  the  morphology  of  the  groups  next  below,  which 
are  themselves  teleological  modifications  of  it ;  conversely,  each  lower  group  is,  with 
reference  to  the  next  higher,  more  directly  teleological,  and  increasingly  so  as  we  ap- 
proach the  species  and  the  individual.  Even  the  types,  ideal  and  unsubstantial  as  they 
seem,  represent  the  four  ways  in  which  the  powers  of  sensation  and  voluntary  motion 
may  be  embodied  and  brought  into  use  in  the  economy  of  Nature  :  the  idea  of  an  animal 
is  distinct  enough  in  our  minds,  but  so  hard  to  put  into  words  that  no  really  satisfactory 
definition  has  ever  been  proposed.  What  better  evidence  of  the  immaterial  character  ot 
the  principle  which  distinguishes  the  animal  from  the  vegetable  and  mineral  subdivisions 
of  Nature. 

It  seems  at  first  rather  strange  that  the  progress  in  philosophical  anatomy  may  be  esti- 


IN  THE   LIMBS  OF   MAMMALIA.  5 

mated  by  the  more  frequent  occurrence  of  the  term  homology,  especially  in  the  works 
of  Professor  Owen,  who  has  done  so  much  toward  dispelling  the  mystery  and  almost 
odium  attached  to  the  subject,  and  has  cleared  up  some  of  its  most  difficult  problems,1 
while  the  philosophy  of  botany  is  measured  by  the  term  morphology,  although  it  has  appar- 
ently never  been  perceived  that  they  are  corresponding  terms,  the  one  relative,  the  other 
absolute. 

But  much  of  the  wonder  vanishes  when  it  is  remembered  that  the  unit  of  vegetable 
structure  is  very  simple,  consisting,  in  the  Dicotyledones,  of  the  phyton,  or  leaf  with  its 
segment  of  stem  ;  and  that  out  of  these,  by  wonderful  transmutation  and  combination,  the 
whole  plant  is  built  up.  The  morphology  of  a  vegetable  organ  is  enunciated  when  it  is 
shown  in  what  manner  it  is  referable  to  the  typical  phyton;  and  since  so  few  elements 
compose  this,  seldom  would  there  arise  questions  of  special  and  thus  of  general  or  serial 
homology.  But  with  animals  the  case  is  otherwise.  Having  left  the  simple  cell,  of  which 
vegetables  also  are  composed,  we  find  at  once  that  their  bodies  are  made  up  of  many  organs 
which  cannot  possibly  be  referred  to  any  one  unit  of  structure.  The  nervous,  circulatory, 
and  digestive  systems  are  entirely  isolated  from  each  other,  and  differ,  not  only  physiologi- 
cally, but  microscopically  and  chemically.  Still  more  complex  are  the  relations  existing 
in  the  muscular  and  osseous  systems,  as  presented  in  the  Vertebrates ;  for  here  the  skele- 
ton is  made  up  of  a  series  of  segments  called  vertebrae,  which  are  themselves  composed 
of  smaller  parts  or  elements  having  definite  relations  and  bearing  distinct  names,  and  by 
variation  in  the  number,  size,  and  shape  of  which  an  almost  endless  diversity  is  produced. 
And  now  the  questions  which  arise  are  emphatically  those  of  relation,  of  homology :  what 
parts  represent  each  other  in  different  animals ;  what  position  one  element  of  a  vertebra 
holds  with  reference  to  the  others  in  the  same ;  and  what  elements  in  two  different  verte- 
bras repeat  each  other  ;  —  questions  of  special,  of  general,  and  of  serial  homology,  respec- 
tively. It  is  not,  then,  so  strange  that  botanists  have  used  the  absolute  term  morphology  with 
reference  to  the  objects  of  their  study,  when  so  few  parts  or  elements  compose  the  morph 
or  type  of  which  the  members  of  any  one  large  group  are  built  up,  as  that  the  anatomist, 
in  his  anxiety  to  determine  the  manifold  relations  existing  in  the  bodies  of  animals, 
should  look  upon  morphology  only  as  the  necessary  guide  to  the  more  difficult  questions 
of  homology,  which  in  itself  implies  more  than  one  morph. 

Teleological  diversities  are  as  of  more  and  less,  and  the  resulting  varieties  communicate 
with  each  other  only  by  continuity ;  by  continuous  degrees. 

Morphological  diversities  are  as  of  interior  and  exterior,  as  of  superior  and  inferior,  and  the 
resulting  varieties  communicate  only  by  contiguity  ;  by  discrete  degrees. 

Here,  if  rightly  appreciated,  is  contained  the  essence  of  two  most  interesting  and  not 
always  easily  understood  generalizations,  which  are  potent  weapons  of  the  modern  zoolo- 
gist :  the  one  defensive  of  his  belief  in  a  natural  classification,  the  other  offensive  against 
those  who  assert  the  existence  of  a  regular,  uninterrupted  succession  of  organic  forms  from 
lowest  to  highest,  because,  forsooth,  they  cannot  see  how  else  creation  was  effected ;  thus 
profanely  daring  to  limit  Infinite  power  by  their  own  wilfully  diminished  capacity. 

1  In  his  elaborate  and   admirable  Report  on  the  Homolo-  another  vertebra  before  or  behind,  its  serial  homology  is  given. 

yies  of  the  Vertebrate  Skeleton,  Professor  Owen  defines  three  3d.    When  a  part  is  said  to  have  the  same  relations  in  two 

relations  of  homology:  "1st.  When  a  part  is  said  to  occu-  different  animals,  then  its  special  homology  is  indicated." 
py  a  certain  position  in  its  vertebra,  its  general  homology  is         These  definitions,  as  we  shall  see,  do  not  cover  all  relations 

enunciated.     2d.    When  such  a  part  is  said  to  repeat  in  its  of  homology. 
vertebra  that   which  occupies   a  corresponding  position   in 
2 


6  WILDER  ON  MORPHOLOGY  AND  TELEOLOGY 

These  are,  1st,  the  law  of  Parallel  Relations ;  and,  2d,  the  teleological  rising  above  or 
sinking  below  their  morphological  level,  of  certain  groups  or  species  or  individuals,  whereby 
they  seem  to  be  of  a  higher  or  lower  grade  than  the  rest  of  the  group  of  which  they  are 
generally  the  extreme  aberrant  forms. 

Illustrations  of  either  of  these  laws  are  almost  superfluous ;  of  the  former  many  will  have 
occurred  to  the  naturalist  who  observes  similar  functions  exercised  by  animals  belonging  to 
different  groups,  or  even  types:  —  the  aerial  bird  and  butterfly;  the  heavy,  graminivorous 
cattle  among  Mammalia,  and  the  phytophagous  Scarabei  among  Coleoptera ;  the  monkey 
and  the  parrot;  the  whole  type  of  Articulates,  and  the  vertebrate  class  Aves;  the  type  of 
Mollusks,  and  the  vertebrate  class  Reptilia ;  the  three  classes  of  Articulates,  with  the  three 
orders  of  its  highest  class,  Insecta ;  and,  finally,  the  striking  parallelism  between  the  orders 
of  the  two  groups  of  Mammalia,  called  by  Dana  Megasthenes  and  Microsthenes,  (American 
Journal  of  Science  and  Art,  vol.  xxxv.  p.  70,)  with  the  less  evident  one  between  the  Altrices 
and  the  Precoces  among  birds.  Between  all  these  pairs  of  groups  is  so  evident  a  similarity 
as  to  have  suggested  the  term  "Parallel  Relation;"  but  it  is  to  be  observed  that  the  rela- 
tion is  one  of  analogy,  not  hoinology ;  that  the  differences  are  morphological,  and  the  resem- 
blances are  comparatively  teleological,  while  between  component  parts  of  the  same  group 
the  resemblances  are  morphological  and  the  differences  teleological. 

Many  insects  are  physiologically  more  highly  organized  than  the  lowest  fishes,  and  the 
eagle  seems  a  creature  vastly  superior  to  the  whale;  but  in  each  case  the  groups  to  which, 
according  to  their  essential  structure,  the  insect  and  the  eagle  belong,  are,  as  groups,  on  a 
plane  below  the  fishes  and  the  mammals.  The  two  relations  are  commonly  expressed  by 
representing  the  groups  by  parallel  vertical  lines ;  there  may  be  such  morphological  differ- 
ences between  the  groups  as  to  clearly  indicate  which  are  higher  and  which  lower,  but  the 
lines  may  be  overlapped,  to  show  that  the  lowest  in  one  group  is  teleologically  inferior  to 
the  highest  of  the  next  group,  though,  as  said  above,  there  would  be  no  doubt  concerning 
the  groups  taken  as  wholes. 

There  is  not,  at  least  among  the  higher  groups,  any  such  lineal  shading  off  into  each 
other  as  to  afford  any  support  to  the  idea  of  a  regular,  uninterrupted  succession  of  organic 
forms,  whether  zoological  or  genealogical.  Nor  does  the  present  state  of  Paleontology  fur- 
nish the  disciples  of  Darwin  much  assistance  in  this  respect. 

Position  may  determine  a  morphology  in  addition  to  that  dependent  upon  structure,  and 
nowhere  is  this  more  clear  than  with  the  teeth  of  Mammalia.  Professor  Owen,  in  his 
Odontography,  has  shown  that  every  classification  of  these  organs  based  upon  their  form, 
and  thus  upon  their  special  masticatory  function,  utterly  fails  in  precision  on  general  appli- 
cation, and  that  the  position  of  the  teeth  in  the  jaws  is  the  only  safe  guide  to  their  arrange- 
ment. In  this  case,  it  so  happens  that  the  teeth  were  originally  named,  from  their  shape 
and  function,  incisors,  canines,  and  molars  ;  and  this  is  the  order  in  which  they  stand  in  the 
jaws  from  before  backward.  But,  while  this  would  answer  very  well  in  designating  the 
corresponding  teeth  in  two  animals  having  the  same  number,  and  where  the  variations  in 
form  were  slight,  it  utterly  failed,  even  in  the  hands  of  Cuvier,  accurately  to  determine 
such  correspondence  when  applied  to  the  whole  range  of  the  mammalian  series. 

Without  entering  into  details,  which  are  given  in  abundance  in  the  Odontography,  it 
may  be  said  that  the  teeth  collectively  are  distinguished  from  all  other  parts  and  organs, 
hard  or  soft,  by  a  peculiar  structure  or  morphology  of  their  own ;  but  that,  to  ascertain  the 
limits  of  the  several  groups  of  tee£h  in  the  jaws  of  a  single  species,  or  to  point  out  corre- 
sponding or  homologous  teeth  in  animals  having  a  different  number,  their  position  in  the 
jaws  is  the  only  safe  standard,  this  constituting  a  minor  morphology. 


IN  THE  LIMBS   OF  MAMMALIA.  7 

• 

Thus  we  have  teeth  morphologically  canine,  but  telcologically  incisor, — the  outer  pair  of  teeth 
in  the  incisive  row  in  the  lower  jaw  of  the  typical  Ruminants,  (sheep,  cattle,  &c. ;)  and,  on 
the  other  hand,  we  have  teeth  which  are  morphologically  incisor,  but  ideologically  canine, — the 
only  pair  of  teeth  in  the  intermaxillary  bone  of  the  Camelidoe,  in  which  aberrant  group  of 
Ruminants  the  lower  canines  above  mentioned  assume  their  proper  form  and  function, 
as  if  to  compensate  for  the  absence  of  horns. 

And  this  brings  us  at  once  to  the  consideration  of  the  important  question,  whether  every 
anatomical  generalization  is  not  an  expression  of  morphology ;  whether  every  grouping  of 
facts  which  we  regard  as  natural,  and  which  enables  us  better  to  comprehend  and  arrange 
other  facts,  is  not  morphology  in  the  strictest  sense  of  the  word.  If  so,  as  I  believe,  then 
all  anatomy  is  or  should  be  morphology ;  for  all  particulars  should  be  studied  with  refer- 
ence to  generals  already  ascertained  or  to  be  elucidated.  And  thus  morphology  comes  to 
be  a  very  simple  thing,  and  not  at  all  a  mystery,  and  will  be  avoided  only  by  those  who 
confound  rational  philosophy  with  unprofitable  imaginings  of  pretty,  pleasing  fancies.  The 
Creator  did  not  work  with  barren  isolated  facts;  and  only  those  who  strive  to  rise  above 
these,  will  ever  gain  an  insight  into  the  way  in  which  He  did  work,  with  general  laws  first 
established,  but  only  with  reference  to  the  particular  ultimate  facts  which  were  grouped 
around  them. 

In  the  human  body  must  exist  just  such  complication  of  structure  and  arrangement  of 
parts  as  best  adapt  it  to  be  the  fit  and  willing  agent  of  the  human  mind ;  and  as  this  is,  if 
not  always  actually,  yet  potentially,  on  a  plane  superior  to  that  of  brutes,  so  we  are  pre- 
pared to  find  in  its  fleshy  covering  a  perfection  of  structure  and  harmonious  arrangement 
of  parts,  which,  in  their  totality,  far  surpass  what  we  observe  in  inferior  animals. 

In  animals,  it  is  true,  there  may  often  exist  a  higher  development  of  one  function  or 
class  of  functions;  but  this,  as  we  shall  see,  is  always  at  the  expense  of  the  rest,  besides  mar- 
ring that  beauty  of  proportion  which  is  really  an  important  element  in  the  human  frame. 
The  fish  and  the  whale  swim  better  than  man,  but  the  form  and  structure  requisite  for  this 
simplest  mode  of  locomotion  render  every  other  impossible ;  even  the  limbs  of  the  seal, 
though  rather  more  free,  are  awkward  imitations  of  anything  unless  it  be  paddles.  The 
teeth  and  stomachs  of  the  strictly  carnivorous  or  herbivorous  animals  are  better  adapted 
for  seizing  and  lacerating  or  chewing,  and  for  digesting  certain  kinds  of  food;  but  the 
necessary  limitation,  as  regards  other  kinds,  is  an  obvious  imperfection,  taking  the  creat- 
ure as  a  wThole.  The  bird  flies  through  the  air  with  a  velocity  which  man  will  probably 
never  equal  by  any  mechanical  contrivance ;  but  the  necessary  concentration  of  weight 
between  the  wings  makes  the  anterior  and  posterior  extremities  mere  bony  supports  for 
air  and  earth,  the  head  taking  the  place  of  the  hand  as  an  organ  of  prehension,  and  becom- 
ing thereby  incapable  of  speech  or  expression.  The  great  strength  of  the  ox,  and  the 
speed  of  the  horse  or  of  the  deer,  are  gained  by  such  an  arrangement  of  the  muscles  of  the 
limbs,  and  modification  of  their  bony  frame,  as  almost  to  preclude  any  other  motion  than 
simple  flexion  and  extension  forward  and  backward,  involving  also  the  loss  of  prehensile 
power  in  the  hand.  Even  the  ape,  whose  structure  is  so  perfectly  fitted  for  climbing,  is, 
so  far  as  regards  the  location  of  the  organs  of  prehension  and  of  progression,  a  man 
reversed;1  and  the  power  of  free  rotation  in  the  forearm,  with  the  great  strength  of  the 
fingers,  is  specially  adapted  to  its  peculiar  mode  of  progression,  and  not  to  the  elevated 
uses  which  the  human  hand  performs. 

1  Contributions  to  the  Comparative  Myology  of  the  Chimpanzee,  Boston  Journal  of  Natural  History,  vol.  vii. 


8  WILDER  ON  MORPHOLOGY  AND  TELEOLOGY 

In  short,  looking  merely  at  man's  body,  beside  being  cosmopolitan  and  typically  omniv- 
orous, although  it  has  a  position  and  mode  of  locomotion  peculiar  to  itself,  and  in  which  it 
certainly  is  unrivalled,  it  is  also  endowed  with  the  power  to  assume  with  grace  almost  every 
conceivable  attitude,  and  to  employ  at  will  the  typical  modes  of  locomotion  of  other  Verte- 
brates, such  as  swimming,  crawling,  leaping,  and  climbing ;  and  all  these  the  human  mind 
has  found  means  to  outstrip  in  point  of  speed;  even  the  flying  of  the  bird,  though  probably 
it  can  never  be  equalled  in  rapidity,  has  been  imitated  by  the  aerial  mode  of  locomotion 
contrived  by  the  same  continent  of  man's  essential  superiority. 

But,  leaving  the  mind  wholly  out  of  view,  the  human  body  is  so  constituted  as  not 
only  to  best  execute  its  own  peculiar  movements,  but  also  to  assume  more  readily  than 
the  brutes  some  which  are  peculiar  to  other  species.  In  other  words,  while  endowed  with 
sufficient  strength  and  firmness  for  all  ordinary  occasions,  it  has  at  the  same  time  such 
flexibility  and  independence  of  action  as  to  be  able  to  apply  this  strength  in  many  and 
very  diverse  ways. 

For  clearer  illustration  let  us  contrast  two  extremes,  the  arm  of  man  with  the  fore-leg  of 
the  horse.  The  former  can  do  nearly  anything  and  everything  except  that  of  which  alone 
the  latter  is  capable,  namely,  to  support  and  propel  the  body  on  the  earth  ;  yet  in  the  two 
limbs  are  the  same  joints,  and,  except  in  the  hand,  the  same  bones  and  muscles ;  but  in  the 
quadruped  the  latter  are  short  and  thick,  and  so  disposed  on  the  front  and  back  of  the  limb 
as  to  pull  it  with  great  force  in  those  two  directions,  and  in  no  other ;  while  in  man  they 
are  arranged  evenly  around  the  bony  shafts,  thus  adding  to  the  symmetry  of  the  limb,  as 
well  as  increasing  its  mobility. 

But  the  most  striking  difference  is,  that,  while  the  movements  of  each  segment  of  the 
human  arm  are,  if  necessary,  entirely  independent  of  those  of  the  other  segments,  in  the  horse 
they  are  much  less  so,  and  flexion  or  extension  at  the  elbow  causes  a  mechanical  movement 
at  the  wrist,  and  vice  versa  :  the  independence  of  the  movements  of  these  two  joints  seems 
to  correspond  with  the  degree  of  development  of  the  humeral  condyles  ;  these  processes, 
when  they  exist,  are  situated  just  above  the  insertions  of  the  external  and  internal  lateral 
ligaments  of  the  elbow,  at  which  two  points  there  is  of  course  no  motion  ;  the  condyles  lie 
a  little  above  and  therefore  change  position,  though  very  slightly,  during  movement  at  the 
joint.  The  extensor  muscles  of  the  wrist  and  fingers  arise  from  the  external  condyle,  the 
flexors  from  the  internal,  both  processes  being  very  prominent  in  man ;  but  it  is  evident 
that  when  the  condyles  are  very  small  or  absent,  the  origins  of  the  muscles  must  in  like 
degree  reach  above  or  below  the  centres  of  motion,  and  thus,  with  the  parts  into  which  they 
are  inserted,  be  more  or  less  affected  by  any  movements  at  the  joint.  Now  in  the  horse 
the  condyles  are  almost  wholly  wanting,  the  flexors  of  the  hand  arising  below  the  centre 
of  motion  on  the  inner  side  of  the  humerus,  and  the  extensors  above  the  corresponding 
point  on  the  outer  side ;  when,  therefore,  the  hand  is  flexed,  the  humerus  and  fore-arm, 
are  also  flexed  at  the  elbow,  and  when  the  hand  is  extended,  these  other  segments  are  also 
extended. 

Hold  the  fore-leg  of  a  horse  horizontally  by  the  part  between  the  elbow  and  wrist,  and 
flex  the  hand ;  the  limb  bends  at  the  elbow  also.  Now  if  you  rest  the  limb  in  its  natural 
position  upon  the  earth,  the  obliquity  in  the  direction  of  the  hoof  tends  to  extend  the  hand 
at  the  wrist,  and  thus  to  straighten  the  limb  at  the  elbow,  so  that  the  heavier  you  press 
upon  the  top  of  the  humerus,  the  firmer  the  limb  becomes.  It  is  evident  that  this  would 
be  a  mechanical  aid  in  sustaining  the  weight  of  the  animal,  but  I  have  had  no  opportunity 
to  look  for  a  similar  arrangement  in  the  posterior  extremity.  On  account  of  this  same 


IN   THE   LIMBS    OF   MAMMALIA.  9 

structure,  however,  the  fore-leg  gives  way  suddenly  and  completely  when  the  animal 
stumbles  so  as  to  bear  upon  the  tip  of  the  hoof  and  so  flex  the  hand  at  the  wrist,  for  that 
brings  the  humerus  down  with  it.  In  the  hind-leg  of  the  frog,  which  is  used  for  little  else 
than  leaping,  there  is  a  somewhat  similar  arrangement,  the  great  extensor  of  the  foot  being 
connected  with  that  of  the  leg  by  a  strong  tendinous  band  on  the  inner  side  of  the  knee, 
so  that  extension  of  one  segment  is  mechanically  connected  with  that  of  the  other.  In  the 
quadrumana  and  carnivora  the  condyles  are  present,  though  less  prominent  than  in  man  ; 
and  indeed  the  degree  of  their  development  seems  to  correspond  nearly  with  that  of  the 
clavicle,  both  of  them  being  concerned  in  the  freedom  and  mobility  of  the  anterior  ex- 
tremity. 


The  new  relations  of  morphology  observed  among  the  muscles  of  the  mammalian  limbs 
are  intimately  connected  with  two  other  generalizations  applying  to  these  parts ;  and  these 
again  are  subordinate  to  the  great  anatomical  law  of  "  antero-posterior  symmetry,"  as  it 
has  been  hitherto  called ;  and  since  little  or  nothing  has  been  published  concerning  this  in 
the  form  it  has  of  late  assumed,  it  may  be  well  to  state  the  law  here,  chiefly  according  to  the 
views  of  Professor  Jeffries  Wyman,  by  whom  it  was  suggested  to  me,  and  who,  almost  alone 
in  this  country,  has  devoted  time  to  eliminating,  from  the  indefinite  and  often  extravagant 
and  absurd  shape  in  which  it  was  left  by  Oken,  the  real  truth  of  a  principle  the  most  potent 
and  elevated  of  which  the  vertebrate  body,  considered  by  itself,  is  capable. 

Yet  in  my  opinion  even  this  is  subordinate  to  a  still  higher  law  which  pertains  as  well 
to  the  other  types  of  the  animal  kingdom,  and  also  coincides  with  a  geometrical  law  so 
closely  as  to  afford  new  ground  for  its  belief.  In  order  to  appreciate  the  full  force  and 
value  of  lesser  laws,  it  must  first  be  shown  how  they  depend  upon  greater  ones ;  and  there- 
fore the  latter  shall  be  first  considered. 

A  partial  statement  of  this  higher  law,  which  for  reasons  given  further  on  I  have  called 
the  law  of  animal  polarity,  was  made  by  Professor  Agassiz,  at  a  meeting  of  the  Boston  Soci- 
ety of  Natural  History,  December  4th,  1861. 

He  characterized  the  four  leading  types  of  the  animal  kingdom  by  four  terms  indicative 
of  the  general  arrangement  of  their  organs,  or  their  plan  of  structure  :  the  Radiates  by 
"  radiality"  the  Mollusks  by  "  laterality"  the  Articulates  by  "  tergality"  and  the  Vertebrates 
by  "  cephality"  In  the  Radiates  all  the  parts  are  disposed  about  a  common  centre,  encircling 
which  also  is  the  dynamic  portion  of  the  nervous  system,  a  ganglion  for  each  diverging 
segment  or  spheromere.  These  spheromeres  are  morphologically  exact  repetitions  of  each 
other,  though  their  size  and  shape  may  be  greatly  modified,  and  even  one  of  them  may  be 
entirely  wanting,  so  that  the  animal  appears  as  if  divisible  into  two  lateral  halves,  when 
really  this  is  due  to  a  teleological  modification  not  at  all  affecting  the  real  plan  of  structure, 
but  only  foreshadowing,  as  it  were,  the  characteristic  arrangement  of  the  next  higher  type, 
just  as  the  molluscan  Bryozoa  present  an  appearance  of  radiation  in  the  disposition  of 
their  groups  of  tentacles.  These  two  instances  show  the  importance  of  always  looking  first 
at  the  more  essential  parts  of  the  body,  rather  than  at  the  outside,  which,  like  other  ap- 
pearances, is  often  deceitful. 

The  laterality  which  Professor  Agassiz  considers  characteristic  of  the  Mollusks,  must 
be  carefully  distinguished  from  the  bilaterality  or  bilateral  symmetry  of  all  animals  above 
Radiates  :  for  the  latter  terms  mean  only  that  the  body  is  composed  of  two  lateral  halves 
which  are  right  and  left  repetitions  of  each  other ;  and  this  is  often  more  conspicuous  in 

3 


10  WILDER  ON  MORPHOLOGY  AND  TELEOLOGY 

the  Vertebrates,  and  especially  in  the  Articulates  with  their  sharply  defined  outlines,  than 
in  the  almost  amorphous  forms  of  the  Mollusca ;  but  here  again  we  must  go  beneath  the 
surface,  and  then  we  find  that  in  the  Mollusks  not  only  are  the  organs  arranged  upon  the 
two  sides  of  the  body,  but  the  "  weight  of  organization,"  as  Professor  Agassiz  expresses  it, 
is  thrown  upon  the  sides,  which  even  in  common  usage  we  recognize  to  have  superior 
value  over  the  front  and  hind  ends,  or  the  upper  and  lower  edges ;  we  examine  and  figure 
only  the  sides,  and,  except  with  the  Cephalopoda,  their  natural  position  is  such  as  to  exhibit 
prominently  one  of  the  sides.  This  distinction  between  the  Ulaterality  common  to  all  above 
Radiates,  and  the  laterality  proper  to  the  Mollusks,  is  well  set  forth  by  Mr.  N.  S.  Shaler, 
in  the  Proceedings  of  the  Boston  Society  of  Natural  History,  December  4th,  1861. 

With  the  type  of  Articulata,  it  is  not  the  right  and  left  sides  that  we  chiefly  regard  in 
either  a  popular  or  a  scientific  examination,  but  the  upper  and  lower  regions,  which  are,  as 
it  were,  set  off  against  each  other.  We  no  more  think  of  placing  or  viewing  an  insect  on 
its  side  than  a  bivalve  on  its  upper  or  lower  edge,  which  correspond  to  the  tergal  and 
ventral  regions  of  the  Articulate.  This  seems  to  confirm  the  idea  that  the  single  ring 
representing  the  articulate  unit  is  composed  of  an  equal  number  of  segments  above  and 
below  a  horizontally  bisecting  plane,  and  that  the  legs  and  wings  when  they  exist  are 
tergal  and  ventral  repetitions  of  each  other.  But  the  internal  anatomy  is  less  satisfactory, 
at  least  as  now  understood,  and  I  leave  it  to  others,  more  familiar  with  its  details,  to 
determine  whether  this  type,  whose  sharply  defined  outlines  so  clearly  illustrate  the  law, 
has  at  the  same  time  the  most  unsatisfactory  internal  arrangement ;  it  is  certain  that  in 
our  present  state  of  knowledge  the  laterality  of  the  Mollusks  is  more  apparent  than  either 
the  tergality  of  the  Articulates  or  the  cephality  of  the  Vertebrates. 

This  latter  term,  cephality ',  applied  by  Professor  Agassiz  to  the  highest  type,  indicates  the 
extreme  preponderance  in  function  of  one  end  of  the  body ;  which,  at  first  on  the  same  level 
with  the  other  end,  is  gradually  raised,  till  it  attains  the  greatest  possible  elevation  in  the 
erect  position  of  man.  Professor  Dana's  term,  "  cephalization,"  is  indicative  of  this  gradual 
ennobling  of  one  end  of  the  vertebrate  body,  and,  in  man,  of  the  devotion  of  the  arms  and 
hands  to  its  requirements,  a  physiological  return  to  an  allegiance  they  always  owed  the 
head,  to  which,  in  fishes,  they  are  actually  attached. 

In  this  connection  it  may  be  added,  that,  besides  the  overwhelming  evidence  adduced  by 
Professor  Owen  in  support  of  the  view  now  generally  received,  that  the  scapular  arch  is  really 
the  modified  pair  of  ribs  of  the  posterior  or  occipital  cranial  vertebrae,  there  are  other  facts 
which  indicate  that  in  the  early  stages  of  even  the  higher  Vertebrates,  the  shoulders  and 
head  are  much  nearer  together  than  in  their  adult  condition. 

1st.  The  singular  course  of  the  inferior  laryngeal  nerve,  whence  comes  its  name  of  the 
recurrent,  is  inexplicable  on  any  other  than  strictly  morphological  grounds ;  for,  instead  of 
proceeding  by  the  shortest  and  most  direct  route  to  the  larynx  from  its  origin  on  the  pneu- 
mogastric,  it  always  forms  a  loop  around  the  subclavian  artery  on  the  right  side  and  the 
arch  of  the  aorta  on  the  left,  even  in  the  giraffe,  where  its  length  is  several  times  as  many 
feet  as  it  would  be  inches  on  the  ground  of  teleology  alone.  An  account  of  a  case  of  mal- 
formation, which  first  drew  attention  to  this  peculiarity,  was  published  in  the  "  Edinburgh 
Medical  and  Surgical  Journal,"  for  1823,  and  the  same  Journal  for  the  month  of  April,  1826, 
contains  an  account  of  a  similar  case,  with  an  explanation  of  this  apparent  waste  of  nervous 
matter.  Both  of  these  accounts  are  quoted  on  page  379  of  "Power's  Surgical  Anatomy  of 
the  Arteries." 

2d.  Professor  Vrolik,  in  his  work  on  Monstrosities,  "Tabulae  ad  illustrandam  Embryogene- 


IN  THE   LIMBS   OF   MAMMALIA.  H 

sin  Hominis  et  Mammalium,"  figures,  and  briefly  describes,  the  skeleton  of  an  anencephalous 
monster,  in  which  one  arm  appears  attached  to  the  base  of  the  skull,  as  if  by  arrest  of  de- 
velopment, while  the  other  is  in  its  normal  position  on  the  side  of  the  thorax.  With  a  view 
to  ascertain  whether,  at  any  period,  the  shoulders  of  the  mammalian  foetus  are  in  actual 
contact  with  the  cranium,  I  made  careful  examination  of  large  numbers  of  foetal  pigs,  and 
in  the  very  smallest,  just  when  the  limbs  begin  to  protrude  from  the  sides  as  little  fleshy 
buds,  it  is  always  at  some  distance  from  the  head ;  so  that,  in  the  Mammalia  at  least,  the 
fact  of  actual  contact  must  be  regarded  as  doubtful. 

3d.  But  in  most  fishes  they  are  firmly  attached  to  the  cranium,  and  in  the  tadpoles  of 
the  bull-frog  (Rana  pipiens)  I  have  found  the  scapula  closely  connected  with  the  poste- 
rior part  of  the  cranium,  either  by  muscle  or  ligament,  which  elongates  as  development 
proceeds. 

Now  the  three  terms,  laterality,  tergality,  and  cephality,  are  more  or  less  complete  expres- 
sions of  the  arrangement  of  organs  at  the  two  poles  of  the  three  axes  of  a  sphere,  the  lateral, 
the  vertical,  and  the  longitudinal,  one  of  which  is  specially  prominent  in  each  of  the  higher 
types,  Mollusks,  Articulates,  and  Vertebrates,  while  the  Radiates  are  represented  by  the  sim- 
ple sphere  itself,  with  no  one  axis  more  prominent  than  another ;  since  the  members  of  this 
type  are  not  geometrical  figures,  but  organic,  living  beings,  they  must  have  a  structural 
axis  around  which  their  diverging  segments  or  spheromeres  are  arranged ;  but  this  assumes 
such  a  variety  of  directions,  being  reversed  between  the  Polyps  and  the  Acalephs  with  most 
Echinoderms,  and  becoming  horizontal  in  the  Holothurians,  as  to  entirely  negative  the  idea  of 
its  having  any  such  morphological  significance  as  the  axes  of  the  other  three  types.  These 
three  are  the  main  axes  of  a  sphere,  the  only  ones  possible  at  right  angles  with  each  other ; 
and  they  also  correspond  with  the  three  dimensions  of  a  solid,  —  breadth,  thickness,  and 
length,  —  while  the  sphere  may  be  regarded  as  having  no  dimension,  yet  as  capable  of  all. 
(See  also  Professor  Agassiz's  "  Contributions  to  Natural  History  of  the  United  States,"  vol.  iii. 
chap.  ii.  sect.  iv.  on  Morphology  and  Nomenclature,  p.  76.)  This  gives  us  four  plans,  four 
morphs  on  which  the  animal  kingdom  is  built,  and  this  coincides  with  the  number  now  be- 
lieved to  exist. 

A  strong  corroboration  from  a  different  source  is  contained  in  the  views  of  Professor  Arnold 
Guyot,  expressed  in  a  course  of  lectures  delivered  during  the  winter  of  1862,  at  the  Smith- 
sonian Institution,  on  the  "  Unity  of  Plan  in  Animals  and  Plants." 

He  presented,  as  an  indication  of  the  existence  of  no  more  nor  less  than  four  grand  divis- 
ions among  animals,  the  idea  that  the  four  types  represent  the  four  grand  epochs  in  the  life 
of  a  single  animal ;  the  Radiates  are  the  starting-point,  the  germ,  the  simple  cell,  with  life, 
but  this  of  a  low,  indeterminate  character,  and  inhabiting  the  water,  the  lowest  medium ; 
then  comes  a  partial  progress  in  one  direction,  with  the  development  of  the  nutritive  sys- 
tems of  organs,  and  this  second  stage  is  represented  by  the  Mollusks,  with  their  heavy  bodies, 
devoted  to  digestion  and  circulation,  and  confined  to  the  earth ;  then  comes  a  partial  prog- 
ress in  the  opposite  direction,  with  the  development  of  the  respiratory  and  motory  apparatus, 
and  this  is  well  represented  by  the  Articulates,  chiefly  inhabiting  the  air ;  and,  finally,  in  the 
Vertebrates  is  typified  the  animal  in  its  perfect  state,  with  a  more  equal  combination  of 
both  classes  of  functions. 

Again,  for  the  existence  of  four  classes  in  the  Vertebrates  the  same  reason  holds  good : 
the  Fishes  are  the  starting-point,  and,  like  the  Radiates,  dwell  in  the  water ;  then  come  the 
Reptiles,  with  their  heavy  bodies  attached  to  the  earth,  and  characterized  by  special  promi- 
nence of  the  nutritive  functions,  thus  corresponding  to  the  Mollusks ;  then  the  Birds,  the 


12  WILDER  OX  MORPHOLOGY  AND  TELEOLOGY 

aerial  Vertebrates,  their  very  bones  filled  with  the  medium  in  which  they  dwell,  and  always 
in  active  motion ;  and  the  cycle  is  again  completed  by  the  Mammals,  which  embody  a  more 
equal  and  harmonious  combination  of  all  the  systems  of  organs,  living,  like  the  Reptiles, 
on  the  earth,  but  elevated  above  it  into  the  free  space  of  air. 

Professor  Guyot  considered  the  same  law  to  prevail  throughout  all  the  lesser  groups 
of  Mammalia,  and  also  in  the  three  other  types ;  and  it  is  certainly  worthy  of  remark,  that 
in  these  latter  are  recognized  only  three  subdivisions,  while  in  the  Vertebrates  four  are  gen- 
erally acknowledged  ;  and  so  in  every  group  containing  man  would  be  four  lesser  groups, 
but  in  all  others  only  three,  the  highest  being  wanting,  and  the  series  thus  incomplete. 

In  this  connection  circle  seems  as  proper  as  series ;  for  the  most  natural  exhibition  of  the 
relative  standing  of  the  four  types  is  by  four  equidistant  points  on  the  circumference  of  a 
circle,  the  Radiates  below,  the  Mollusks  above  and  to  the  left,  the  Articulates  the  same 
distance  above  and  to  the  right,  and  the  Vertebrates  at  the  top  ;  the  two  intermediate 
groups,  the  types  of  partial  progress,  being  at  the  same  distance  from  the  lowest  and 
highest,  and  thus  of  equal  rank,  though  opposite  nature.  Nor  can  we  overlook  the  fact  that 
there  are  four  regions  of  the  body,  pelvis,  abdomen,  thorax,  and  head  ;  and  that,  as  I  think 
Oken  has  said,  Fishes  are  pelvic  animals,  Reptiles  abdominal,  Birds  thoracic.,  and  Vertebrates 
cephalic  ;  and  that  neither  Radiates  nor  Mollusks  nor  Articulates  possess  a  distinct  anterior 
segment  containing  any  such  overruling  portions  of  the  nervous  system  as  does  the  head 
of  the  Vertebrates. 

There  are  four  senses  also,  one  general,  the  others  special. 

The  sense  of  touch  is  universal,  and  only  more  or  less  developed  in  different  regions  of 
the  body  :  it  is  most  exquisitely  perceived  through  the  agency  of  water  or  moisture, 
especially  in  the  tongue  ;  for  taste  is  a  peculiar  exaltation  of  the  general  sensibility,  and 
forms,  as  it  were,  a  transition  therefrom  to  the  smell,  one  of  the  three  special  senses  with 
which,  as  anatomy  clearly  shows,  it  cannot  be  allied.  Of  these  latter,  smell  is  the  lowest, 
and  its  exercise  depends  upon  the  presence  of  odoriferous  particles  of  an  earthy,  solid 
nature.  Hearing  is  the  second  special  sense,  and  perceives  vibrations  in  the  atmosphere, 
the  next  higher  medium ;  while  sight,  the  highest,  depends  for  the  exercise  of  its  functions, 
not  upon  this,  but  upon  that  invisible  and  imponderable  yet  material  medium  which  is 
called  ether. 

Again,  sight  is  directly  related  to  the  central  nervous  system,  and  properly  belongs  to  the 
head,  below  which  its  organ  does  not  extend.  Hearing  goes  lower  down,  into  the  -fhwrynx, 
with  which  its  organ  is  connected,  anatomically  by  the  Eustachian  tube  and  physiologically 
when  we  listen  to  the  speech  of  another.  Smell  and  its  accessory  taste  preside  over  the 
entrance  to  the  alimentary  canal,  with  which  they  descend  through  the  head  and  thorax 
into  the  abdomen.  Finally,  in  the  pelvis  is  the  organ  which,  as  will  be  hereafter  shown,  is 
the  posterior  or  reversed  repetition  of  the  tongue,  and  whose  sensitiveness,  like  taste,  is  only 
a  peculiar  exaltation  of  the  universal  sense,  the  nerves  in  both  cases  being  the  common 
cranial  or  spinal  nerves,  and  not  special  prolongations  of  the  brain  into  the  organ,  as  with 
the  eye,  the  ear,  and  the  nose. 

Professor  James  D.  Dana,  in  a  paper  "  On  the  higher  Subdivisions  in  the  Classification  of 
Mammalia,"  (American  Journal  of  Science  and  Arts,  vol.  xxxv.  January,  1863,)  proposes  a 
similar  quaternary  division  of  that  class  with  special  reference  to  the  "  cephalization"  of 
the  body,  which  he  shows  to  increase  as  we  ascend  in  the  scale. 

To  those  who  make  classification  their  study  it  belongs  to  decide  how  much  influence 
these  principles  exert  among  the  lower  groups ;  but  certainly  among  the  higher  ones  the 


IN  THE  LIMBS   OF   MAMMALIA. 


13 


coincidences  are  too  striking  to  be  disregarded  by  the  most  matter-of-fact  philosopher.     I 
subjoin  a  diagram  exhibiting  the  more  evident  correspondences  in  the  various  departments 


i 


bo 

.5 


O 


1 

4 


o 


PH 


14  WILDER  ON  MOPHROLOGY  AND  TELEOLOGY 

of  Nature,  and  which  seem  to  indicate  that  here  is  one  and  the  same  thought  of  the  Creator 
expressed  in  terms  geometrical,  morphological,  physiological,  and  zoological. 

From  what  has  been  said  concerning  the  axes  of  the  three  higher  types,  it  will  appear 
that  they  all  exist  in  each  of  the  types,  just  as  they  would  in  a  geometrical  sphere,  variously 
compressed  ;  but  that  this  presence  of  all  must  be  carefully  distinguished  from  the  prominence 
of  one  axis  in  the  type  characterized  thereby.  Thus  the  laterally  of  Mollusks  is  something 
more  than  the  lilaterality  or  bilateral  symmetry  which  exists  also  in  the  Articulates  and 
Vertebrates ;  the  cephality,  or  more  properly  the  longitudinality  of  the  latter  typ«  must  be 
distinguished  from  the  antero-posterior  symmetry  so  evident  in  certain  Articulates ;  and  the 
tergality,  or  more  properly  verticality  of  these,  from  any  resemblance,  if  it  should  be  observed, 
between  the  dorsal  and  ventral  regions  of  the  Mollusca ;  even  the  radiality  of  the  lowest 
type  must  not  be  confounded, with  the  appearance  of  it  in  certain  Mollusks  above  mentioned. 

But,  in  fact,  mere  resemblance  between  any  two  regions  of  the  body  is  not  what  we  desire 
to  express ;  for,  although  it  may  exist,  there  is  quite  as  often  a  striking  contrast,  as  in  the 
Gasteropoda  among  Mollusks,  and  in  Man  with  most  Mammalia  among  Vertebrates.  Our 
term  should  refer  rather  to  the  direction  of  the  axis  which  is  specially  prominent  in  any 
one  type,  at  the  poles  of  which  are  the  two  regions  which  are  the  external  evidence  of  this 
prominence,  and  which  morphologically  repeat  each  other,  but  as  has  been  instanced,  may 
be  teleologically  most  diverse ;  and  therefore  if  radiality  and  laterally  are  accepted  for  the 
two  lower  types,  (for  despite  our  morphological  equality  of  Mollusks  and  Articulates,  the 
latter  are  physiologically  the  higher  of  the  two,)  the  corresponding  terms  for  the  other  types 
are  verticality  and  longitudinality :  for  tergality  and  cephality  only  express  the  prominence  of 
the  region  at  one  end  of  the  axis  over  that  at  the  other,  a  prominence  which  is  teleological, 
while  our  idea  is  strictly  morphological.  Radiality,  laterality,  verticality,  and  longitudinality 
are,  morphologically,  peculiar  to  and  characteristic  of  Radiates,  Mollusks,  Articulates,  and 
Vertebrates  respectively,  but,  teleologically,  they  may  be  as  it  were  ingrafted  upon  the 
others  in  the  shape  of  radiation,  bilaterality,  tergality,  and  antero-posterior  symmetry. 

In  accordance  with  the  law  of  polarity,  we  shall  expect  to  find  the  organs  at  the  two 
poles  of  a  prominent  axis  morphologically  similar,  and  thus  to  observe  relations  of  polar 
homology  between  them, — of  lateral  homology  in  the  Mollusks,  of  vertical  homology  in  the 
Articulates,  and  of  longitudinal  homology  in  the  Vertebrates  ;  all  existing  more  or  less  in 
each  of  these  three,  but  specially  prominent  one  in  each  ;  while  in  the  Radiates  there  is  the 
radial  homology  so  early  recognized  between  the  four  or  five  diverging  segments.  These 
four  kinds  of  polar  homology  are  the  most  general. 

We  come  now  to  consider  more  particularly  that  relation  of  homology  which  is  charac- 
teristic of  the  Vertebrates,  and  included  in  the  idea  of  longitudinality,  otherwise  expressed 
by  the  compound  term  antero-posterior  symmetry ;  both  these  terms  are  objectionable 
on  the  score  of  length,  but  the  former  is  really  significative  of  the  idea  we  mean  to  convey. 

It  may  be  well  at  this  point  to  find  how  fully  our  deductions  are  borne  out  by  the  facts. 

It  is  fortunate  indeed  when  our  views  a  priori  are  confirmed  by  our  investigations  a 
posteriori,  so  that  we  safely  ascend  and  descend  the  hill  of  science,  without  being  dazzled 
and  led  astray  by  glittering  and  apparently  correct  generalizations,  or  losing  our  way 
amidst  the  labyrinth  of  facts,  and,  in  our  eagerness  to  advance,  in  place  of  steadily  looking 
for  the  landmarks  Nature  has  provided,  becoming  impatient  and  exhausting  our  strength 
in  cutting  down  or  rooting  up,  if  possible  to  destroy,  whatever  stands  in  our  way,  or  perhaps 
using  it  to  bridge  over  gaps  in  our  self-made  path  which  never  would  have  existed,  if,  at 
the  outset,  our  aim  had  been  to  discover,  not  to  create,  to  learn  from  God,  not  to  be  teach- 
ers ourselves. 


IN  THE    LIMBS    OF    MAMMALIA.  15 

If  the  previous  reasoning  is  correct,  it  follows  that  the  anterior  and  posterior  regions  of 
the  vertebrate  body,  consequently  of  our  own,  are  anterior  and  posterior  repetitions  of 
each  other,  but  in  opposite  directions,  like  the  right  and  left  sides. 

At  first  sight,  nothing  is  more  improbable  to  our  thought  or  revolting  to  our  feelings,  and 
the  evidence  must  be  strong  to  overcome  this  natural  repugnance  to  the  relationship. 

Yet  why  should  it  be  objectionable,  any  more  than  that  neither  anatomy  nor  microscopy 
nor  chemistry  can  detect  the  slightest  difference  between  two  human  brains  to  indicate 
that  one  of  them  was  the  agent  of  a  wicked  and  depraved,  the  other  of  a  noble  and  lofty 
soul  ?  It  is  the  universal  distinction  between  morphology  and  teleology,  between  the  thing 
as  it  is  made  and  as  it  is  used ;  and  when  this  distinction  is  once  rightly  appreciated,  we 
shall  be  no  more  shocked  at  the  morphological  identity  of  the  two  ends  of  our  corporeal 
frame  than  we  are,  or  ought  to  be,  at  the  close  anatomical  relationship  between  the  disgust- 
ing ape  and  ourselves,  remembering  in  both  cases  that  it  is  the  use  alone  which  can  ennoble 
or  debase. 

Our  evidence  is  as  yet  by  no  means  complete  as  regards  all  the  organs  of  the  vertebrate 
body,  but  enough  has  been  brought  out  to  very  strongly  confirm  the  principle. 

As  was  implied  when  treating  of  the  general  law  of  polarity,  we  must,  if  this  is  acknowl- 
edged, recognize  a  new  relation  of  homology  between  parts  representing  each  other  at  the 
two  ends  of  the  longitudinal  axis,  and,  in  fact,  on  opposite  sides  of  the  longitudinal  centre, 
similar  to,  but  by  no  means  so  obvious  as,  that  existing  between  the  right  and  left  sides. 

This  is  not  the  "  serial  homology,"  of  Professor  Owen,  for  he  regarded  all  the  vertebrse  and 
their  appendages  as  simple  repetitions  of  each  other  from  one  end  of  the  body  to  the  other, 
without  recognizing  the  fact,  which  we  shall  see  most  clearly  in  the  limbs,  that  the  anterior 
and  posterior  organs  of  the  body  are  repetitions  of  each  other,  but  in  opposite  directions ;  so 
that  the  term  "  homotype,"  by  which  he  designated  parts  serially  homologous,  really  applies 
only  to  such  as  repeat  each  other  on  one  and  the  same  side  of  the  longitudinal  centre,  and 
for  the  new  relation  between  parts  on  opposite  sides  of  this  point  a  new  term  must  be 
found. 

Adopting  the  phraseology  of  Professor  Owen  we  may  call  all  parts  which  repeat  each  other 
as  opposite  ends  of  any  axis  "  antitypes,"  and  the  antitypes  of  the  lateral  axis  "  latitypes," 
of  the  vertical  axis  "  vertitypes,"  and  of  the  longitudinal  axis  "  longitypes,"  which  will  be 
specially  characteristic  of  the  three  higher  sub-kingdoms,  Mollusks,  Articulates,  and  Verte- 
brates, respectively,  while  the  homologous  diverging  segments  of  the  Radiata  may  be  called 
"  raditypes."  In  the  Vertebrates  the  head  and  pelvis  are  longitypes ;  and  the  thorax  and 
abdomen  bear  the  same  relation.  This  is  a  very  general  homology. 

There  will  now  arise,  with  reference  to  the  longitypes,  or  the  homologous  parts  in  the 
anterior  and  posterior  regions  of  one  animal,  a  question  similar  to  that  concerning  corre- 
sponding parts  in  two  different  species ;  which  was  whether  they  can  be  to  the  same  extent 
homologous  in  animals  bearing  different  degrees  of  zoological  relationship:  for  example,  the 
anterior  extremities  of  all  Vertebrates  are  homologous ;  but  surely  the  arm  of  the  monkey 
is  more  closely  related  to  the  fore-leg  of  the  cat  than  either  is  to  the  flipper  of  the  porpoise 
or  the  pectoral  fin  of  the  fish ;  and  now,  since  the  main  axis  or  vertebral  column  begins 
to  be  formed  at  what  is  afterward  the  point  of  division  between  the  fore  and  hind  regions 
of  the  body,  and  the  head  and  pelvis  are  situated  at  or  near  the  ends  of  that  axis,  it  does 
not  seem  possible  that  these  parts  can  be  as  strictly  homologous  in  animals  having  a 
different  number  of  vertebras  as  in  those  with  the  same  number :  in  other  words,  the  heads 
or  the  pelves  of  two  animals  may  be  cranial  or  pelvic  modifications  of  vertebrse  without 
being  such  modifications  of  the  same  identical  vertebrse. 


16  WILDER   ON    MORPHOLOGY   AND   TELEOLOGY 

We  have  seen  above  that  the  vertebrate  body  is  composed  of  four  regions- — head,  thorax, 
abdomen,  and  pelvis.  The  tail  is  a  pelvic  prolongation,  and  not  a  distinct  region,  for  though 
in  the  lowest  class  it  constitutes  so  large  a  part  of  the  whole  body,  and  is  the  chief  agent  in 
locomotion,  yet  it  contains  no  viscera,  becomes  more  and  more  abbreviated,  and  at  last  dis- 
appears as  we  ascend,  or,  according  to  Prof.  Dana,  as  the  vertebrates  become  "  cephalized." 
The  number  four  at  once  furnishes  a  basis  for  a  twofold  longitudinal  division,  the  head 
and  thorax  in  front,  the  abdomen  and  pelvis  behind ;  the  head  and  pelvis  are  the  extreme, 
and  the  thorax  and  abdomen  the  intermediate,  antero-posterior  representatives  or  anti- 
types of  each  other.  Superficially  or  physiologically,  the  thorax  seems  better  to  repeat  the 
pelvis ;  but  this  is  due  to  the  fact  that  in  most  vertebrates  the  anterior  extremities  are 
shifted  back  upon  the  sides  of  the  thorax  :  which,  of  course,  cannot  be  of  importance  in  a 
morphological  point  of  view ;  so  that,  so  far  as  the  limbs  are  concerned,  and  we  shall  find 
in  them  the  law  of  longitudinality  beautifully  carried  out,  the  head  is  more  clearly  shown 
to  be  the  anterior  representative  of  the  pelvis.  It  is,  of  course,  assumed  that  the  verte- 
bral theory  of  the  skull  is  true,  and  that  it  also  applies  to  the  other  or  caudal  extremity  of 
the  vertebral  column,  where  the  physiological  degradation  is  usually  as  great  as  is  the 
elevation  anteriorly ;  with  the  difference  that  the  tendency  to  linear  multiplication  is 
usually  limited  to  four  cranial  vertebrae,  while  the  tail  may  vary  greatly  in  the  number  of 
its  aborted  vertebral  segments.  The  four  regions  of  the  body  are  associated  physiologically 
also  in  the  same  way  :  the  two  extremes  are  regions  of  relation  ;  their  functions,  sexual  and 
mental,  are  exercised  with  direct  reference  to  other  individuals,  and  the  latter  is  capable  of 
elevation  to  the  highest  communion  possible  between  the  finite  and  the  infinite.  But  the 
functions  of  the  two  intermediate  regions  are  of  a  personal  and  selfish  nature,  those  of  the 
abdomen  being  concerned  in  the  support  of  the  life  of  the  individual  within  itself,  and  those 
of  the  thorax  evolving  the  power  necessary  for  motion  and  for  influence  upon  the  world. 

The  question  next  arises,  Where  is  the  longitudinal  centre  of  the  body  ?  This  cannot  be 
indicated  precisely,  and  perhaps  varies  in  different  species,  but  it  undoubtedly  lies  between 
the  two  intermediate  regions,  thorax  and  abdomen,  perhaps  at  that  vertebra  whose  spinous 
process  is  upright,  inclining  neither  backward  like  those  of  the  dorsal,  nor  forward  like 
those  of  the  lumbar  vertebras ;  but  this  point  will  seem  to  shift  its  position  according  to 
the  various  physiological  requirements  as  to  the  length  of  the  thorax  or  abdomen,  or  the 
strength  and  mobility  of  the  extremities. 

Questions  of  morphology  are  often  determined  by  embryology ;  —  by  reference  to  the 
early  stages  of  development  before  teleological  modifications  have  been  superadded ;  the 
development  of  the  vertebral  column  and  of  the  enclosed  myelon,  conclusively  shows  that 
the  morphological  centre  of  the  body  is  at  the  middle  of  the  back,  notwithstanding  the 
cephalic  end  afterward  acquires  such  physiological  superiority.  Ossification  of  the  vertebral 
column  commences  at  the  middle  of  its  length,  and  proceeds  forward  and  backward  from 
this  point :  the  size  of  the  bodies,  or  centra,  of  the  vertebras  at  this  early  period,  diminishes 
anteriorly  and  posteriorly,  and  only  later  becomes  more  equal,  which  latter  proportion 
persists  through  life  in  the  lower  mammalia  but  in  the  quadrumana  and  especially  in  man 
the  posterior  centra  become  thicker  and  stronger,  so  as  to  give  the  column  that  slender 
pyramidal  shape  necessary  for  the  preservation  of  their  more  or  less  erect  position,  while 
the  upper  or  neural  arch  is  anteriorly  expanded  to  accommodate  the  enormously  enlarged 
cerebral  ganglia,  the  long  axis  of  which  forms  a  decided  angle  with  that  of  the  cord. 

Perhaps  nowhere  is  the  distinction  between  morphology  and  teleology  more  evident  than 
in  the  cerebro  spinal  axis.  At  the  first  appearance  of  this  in  the  embryo,  there  is  no 


IN   THE   LIMBS    OF  MAMMALIA.  17 

such  marked  preponderance  of  the  anterior  portion  as  in  the  later  periods,  but  the  two 
ends  present  a  nearly  similar  appearance,  which  similarity  is  persistent  in  the  lowest  fishes. 
There  seems  to  be  no  good  reason  for  regarding  the  anterior  enlargement  of  the  myelon 
as  essentially  or  morphologically  distinct  from  the  posterior.  The  brain  is  an  after-growth, 
for  teleological  cause,  and  the  posterior  enlargement  is  distinct,  and,  though  entirely  over- 
balanced by  the  immense  development  of  the  organs  of  the  mind  and  special  senses,  is  of 
no  small  importance  even  physiologically.  No  one  who  has  suffered  the  excruciating  pain 
in  the  small  of  the  back,  accompanying  most  febrile  diseases,  will  question  the  importance  of 
the  portion  of  the  myelon  there  situated,  to  which  part  also  are  referred  the  sensations 
of  relief,  more  or  less  distinctly  felt,  upon  the  discharge  of  the  contents  of  intestine,  blad- 
der, uterus  or  testis. 

To  avoid  misconception,  it  may  here  be  stated  that  from  this  morphological  point  of 
view,  I  consider  the  brain  proper  as  a  purely  physiological  addition  with  no  posterior  repre- 
sentative, and  the  so-called  anterior  and  posterior  enlargements  of  the  spinal  cord  as  bun- 
dles of  the  fibrous,  that  is,  connecting  or  adynamic  nervous  substance,  increased  in  size 
and  number ;  but  the  medulla-oblongata  as  the  true  anterior  morphological  enlargement, 
the  posterior  representative  or  longitype  of  which  is  that  apparently  insignificant  portion  of 
the  cord  which  lies  behind  the  origins  of  the  great  lumbar  nerves,  but  which  contains  a 
very  large  proportion  of  the  gray  or  cellular  or  dynamic  nervous  matter. 

But  this  point,  and  many  others,  demand  investigations  more  rigid,  minute,  and  even 
microscopic,  and  with  reference  to  the  principle  involved.  The  idea  of  antero-posterior 
symmetry  is  as  yet  illustrated  to  us  only  by  such  parts  and  organs  as  conform  to  it  mpre 
obviously  by  teleological  likeness  or  unlikeness  in  the  two  regions  of  the  body ;  and,  though 
there  is  already  enough  to  establish  beyond  a  doubt,  to  my  mind  at  least,  the  truth  of  this 
subdivision  of  the  higher  law  of  polarity,  yet  there  is  needed  more  time  and  labor  than 
any  one  has  yet  been  able  to  bestow,  to  demonstrate  the  more  obscure  relations  of  those 
parts  which  are  only  morphologically  anterior  and  posterior  repetitions  of  each  other. 

It  is  by  no  means  unlikely  that  further  investigations,  broad  and  impartial,  may  show 
that,  except  between  certain  organs,  the  relation  of  polar  homology  is  a  general  and  not  a 
special  one ;  at  any  rate,  I  think  we  are  not  yet  prepared  to  state  what  bones  of  the  head 
and  pelvis  are  longitypes ;  if  mere  consolidation  and  organic  connection  of  the  more  an- 
terior vertebrae  constitutes  the  cranium,  then  many  fishes  have  five  or  six,  or  even  more 
cranial  vertebrae,  while  in  the  frog  the  occipital  segment  is  so  divided  that  the  two  lateral 
elements,  or  neurapophyses,  are  separated  from  the  centrum  and  neural  spine,  and  so  appear  to 
constitute  a  distinct  vertebra. 

Perhaps  the  two  or  even  the  three  anterior  cranial  vertebrae,  the  nasal,  frontal,  and 
parietal,  are,  like  the  special  senses,  not  represented  posteriorly.  It  may  be  said  in 
favor  of  this  view,  that,  while  the  universal  sense  of  touch  is  perfected  in 'the  hands,  which 
are  the  distal  ends  of  the  diverging  appendages  of  the  lower  or  haemal  arch  of  the  poste- 
rior or  occipital  cranial  vertebra,  and  taste,  the  modification  of  this  universal  sense,  is 
located  in  the  tongue  which  is  supported  by  the  haemal  arch  of  the  next  or  parietal  verte- 
bra, the  organs  of  the  three  special  senses  are  located  in  or  between  the  superior  or  neural 
arches  of  this  and  the  two  remaining  segments,  which  indeed  seem  to  exist  only  with  ref- 
erence to  these  and  to  the  brain  proper,  none  of  which  have  posterior  representatives ; 
therefore,  we  might  not  expect  to  find  in  the  pelvis  any  parts  corresponding  exactly  to 
these  special  developments,  but  only  the  entire  vertebra  whose  haemal  arch  has  for  its 
diverging  appendages  the  posterior  extremities,  with  that  portion,  hasmapophysial,  of  the 

5 


18  WILDER  ON  MORPHOLOGY  AND  TELEOLOGY 

succeeding  haemal  arch  which  supports  the  penis,  the  longitype  of  the  tongue,  the  pleura- 
pophysial  elements  of  the  arch  being  deficient. 

In  most  mammalia,  but  by  no  means  in  all  vertebrates,  the  pubic  bones  support  the 
male  organ  of  generation;  the  ischiatic  bones  are  certainly  the  posterior  of  the  two  pelvic 
arches,  and  may  reasonably  be  supposed  to  represent  the  hyoid  bone  ;  at  any  rate,  despite 
the  vast  teleological  discrepancy,  the  anterior  portion  of  the  head  seems,  so  far  as  morphol- 
ogy is  concerned,  to  be  the  fixed  longitype  of  the  more  or  less  movable  series  of  vertebras 
behind  the  fixed  pelvis. 

Having  thus  admitted  a  general  homology  between  these  two  extreme  regions  of  the 
body,  we  must  wait  for  further  evidence  to  show  first,  whether  there  is  really  any  such  spe- 
cial homology  as  at  first  appears  probable,  and  if  so,  whether  the  clavicle  and  coracoid  repre- 
sent the  ischium  and  pubis  respectively,  as  the  bones  alone  would  indicate,  or  whether,  regard- 
ing also  the  fleshy  parts,  the  hyoid  bone  is  the  antitype  of  the  ischium,  and  the  clavicle  that 
of  the  pubis,  the  coracoid  being  merely  a  process  like  the  marsupial  bone  of  the  mammals 
characterized  thereby. 

The  alimentary  canal  with  its  appendages  next  merits  our  attention. 

The  former  is  embryologically  and  morphologically  a  simple  straight  tube,  with  an  an- 
terior opening  in  the  head  and  a  posterior  one  in  the  pelvis.  In  the  adult  state,  however, 
this  tube  is  always  more  or  less  enlarged  and  convoluted,  to  afford  a  reservoir  of  the  size, 
and  an  absorbent  surface  of  the  extent  required  by  the  nature  of  the  food  consumed;  but 
all  such  modifications  are  purely  teleological,  and  only  conform  to  the  general  arrangement 
which  assigns  to  the  organs  of  nutrition  a  region  below  that  in  which  are  the  organs  of  the 
motive  forces,  and  above  that  devoted  to  generation. 

There  is  at  first  but  a  single  anterior  and  posterior  opening,  but  the  former  is  generally, 
and  the  latter  in  mammalia  afterward,  divided  into  a  superior  and  an  inferior  opening ;  the 
nasal  and  oral,  the  anal  and  genital  orifices,  by  transverse  bands  which  bear  the  names  of 
upper  lip  and  perinaaum. 

With  one  exception,  all  sexual  diversity  is  teleological,  that  is,  resulting  from  a  difference 
in  the  size  or  shape  of  parts  which  exist  alike  in  both  sexes.  The  exception  is  in  the  case 
of  the  ovary  and  the  testis,  which,  being  entirely  distinct  in  every  respect,  constitute  the 
only  morphological  difference  between  the  two  sexes ;  a  true  hermaphroditism  is,  of  course, 
impossible,  except  between  the  two  sides  of  the  body;  and  this  would  be  a  positive  condi- 
tion, while  all  the  so-called  cases  of  hermaphroditism  are  merely  negative  and  doubtful 
states. 

In  the  anterior  region,  enumerating  from  above,  that  is,  from  the  vertebral  column 
downward,  the  parts  are,  the  nose,  or  anterior  nares,  the  upper  lip,  the  mouth,  the  tongue, 
and  the  chin;  posteriorly,  the  anal  opening,  the  perinceum,  the  vaginal  opening,  the  penis  or 
the  clitoris,  and  the  pules.  The  morphological  correspondence  is  as  evident  as  is  the  tele- 
ological difference. 

There  are  two  principal  diverticula  of  the  alimentary  canal,  the  lungs  and  the  urinary 
bladder;  the  former  open  forward  and  the  latter  opens  backward,  and  their  outlets  are  be- 
tween the  pharynx  or  mouth  and  the  tongue,  anteriorly,  and  between  the  vaginal  open- 
ing and  the  clitoris  posteriorly.  There  is  a  physiological  relation  also,  for  the  bladder  is  a 
dilatation  of  the  internal  portion  of  the  allantois,  which  was  the  foetal  organ  of  respiration. 
The  thyroid  gland  is  in  relation  with  the  larynx  much  as  the  prostate  gland  is  with  the 
neck  of  the  bladder ;  but  the  former  has  no  excretory  duct  as  has  the  latter. 


IN  THE  LIMBS   OF   MAMMALIA.  19 

The  heart,  notwithstanding  its  lofty  physiological  preeminence,  is,  morphologically, 
only  a  more  or  less  complicated  enlargement  and  convolution  of  the  great  arterial  trunk, 
just  as  the  brain  and  stomach  are  teleological  modifications  of  more  simple  fundamental 
parts ;  it  and  the  stomach  are  examples  of  lateral  displacement  from  their  normal  position 
as  median  organs.  Terminating  the  vagina  is  the  uterus,  of  which  the  longitype  is  not 
yet  discovered ;  the  mouth  is  the  longitype  of  the  vaginal  opening  into  the  alimentary 
canal,  of  which  latter,  however,  it  is  the  teleological  inlet.  Pathology  seems  to  indicate 
that  the  testes  and  the  parotid  glands  are  longitudinally  homologous ;  for  inflammation 
of  the  former  is  very  prone  to  invade  the  latter,  by  what  is  called  metastasis,  but  which  in 
this  case  may  be  a  physiological  indication  of  a  morphological  relation  otherwise  obscure. 
So,  likewise,  are  connected  the  diseases  and  their  remedies,  of  the  genito-urinary  and 
respiratory  passages,  and  both  these  cases,  with  that  of  the  irritation  of  the  nostrils  sym- 
pathetic with  the  presence  of  worms  in  the  rectum,  are  simply  analogous  to  what  so  often 
happens  between  parts  which  are  laterally  homologous.  I  am  not  aware  that  any  dispo- 
sition has  yet  been  proposed  of  the  other  abdominal  viscera,  liver,  spleen,  pancreas,  and 
kidneys  ;  —  and  will  merely  refer  to  the  view  of  J.  Maclise  concerning  the  two  former,  that 
they  are  laterally  complementary,  as  stated  on  page  153  of  his  "Surgical  Anatomy,"  where 
are  also  given  several  reasons  for  his  opinion. 

We  come  now  to  the  limbs,  of  which  there  are,  in  vertebrates,  two  pairs:  the  one  anterior, 
and  the  other  posterior.  Their  general  homology  as  diverging  appendages  of  the  haemal 
arches  of  the  occipital  or  posterior  cranial,  and  of  the  anterior  pelvic  vertebrae,  has  been 
already  indicated ;  and  also  the  doubt  as  to  the  general  and  antitypical  relations  of  the 
lower  or  heemapophysial  portions  of  those  arches,  supposed  to  be  represented  by  the  four 
bones,  clavicle  and  coracoid,  ischium  and  pubis.  In  many  mammalia  the  two  former  exist 
only  in  the  shape  of  processes  from  the  scapula,  which  is  the  upper  or  pleurapophysial  por- 
tion of  the  arch;  this  and  its  antitype  the  ilium,  seem  to  follow  the  direction  of  the  devel- 
opment of  the  vertebral  column,  the  former  pointing  forward,  and  the  latter  backward.  To 
the  lower  or  distal  ends  of  these  elements,  by  the  shoulder  and  hip  joints,  are  articulated  the 
proximal  ends  of  the  limbs  proper ;  these  are  made  up  each  of  four  segments ;  anteriorly, 
of  the  arm,  fore-arm,  hand,  (wrist  and  palm,)  and  fingers  ;  or,  osteologically,  of  the  humerus, 
ulna  and  radius,  carpus  and  metacarpus,  and  phalanges ;  and  posteriorly,  of  the  thigh,  leg 
foot,  (ankle  and  instep,)  and  toes ;  or,  of  the  femur,  tibia  and  fibula,  tarsus  and  metatarsus, 
and  phalanges. 

It  is  worthy  of  remark,  that  the  number  of  component  parts  of  the  segments  increases 
toward  the  distal  ends,  while  their  individual  mobility  is  diminished  in  the  same  direction 
as  if  for  mutual  compensation,  and  in  accordance  with  the  general  rule  that  the  right  of  dis- 
cretion increases  and  diminishes  with  responsibility. 

Now  the  law  of  polarity  is  morphological ;  but  in  this  case,  the  teleology  also  is  very  evi- 
dent ;  the  divergence  of  the  scapula  and  ilium  is  on  the  principle  of  a  pyramid,  the  base 
being  wider  than  the  top,  so  as  to  afford  a  firmer  support;  we  shall  find  the  same  polarity 
carried  out  in  the  segments  of  the  limbs  themselves  with  one  exception,  the  necessity  of 
which  is,  however,  as  obvious  as  the  grounds  for  the  general  arrangement  therein  departed 
from;  and  indeed  throughout  the  limbs,  which  as  "a  whole  are  teleological  superadditions, 
the  uses  of  the  general  laws  are  so  apparent  that  the  latter  seem  almost  teleological,  the 
two  principles  being,  as  it  were,  blended  and  thoroughly  harmonious. 

The  general  correspondence  between  the  four  segments  of  the  anterior  and  posterior 


20  WILDER  ON  MORPHOLOGY  AND  TELEOLOGY 

limbs  is  evident ;  the  arm  or  humerus  and  the  thigh  or  femur,  the  fore-arm  and  the  leg, 
the  hand  and  the  foot,  the  fingers  and  the  toes  are  easily  seen  to  occupy  similar  positions 
in  the  limbs  to  which  they  belong ;  but  the  law  of  longitudinality  is  further  carried  out 
among  these  segments  according  to  what  may  be  called  corollaries  thereof. 

1st.  Two  corresponding  segments  in  the  fore  and  hind  limbs  point,  and  are  flexed  or  ex- 
tended, in  absolutely  opposite  though  relatively  similar  directions. 

2d.  Two  contiguous  segments  of  the  same  limb  also  point,  and  are  flexed  or  extended  in 
opposite  directions,  so  that  \hzflexor  muscles  of  one  segment  lie  on  the  same  side  of  the  limb 
with  the  extensors  of  the  segment  next  above  or  below,  and  vice  versa. 

All  this  is  easily  seen  in  the  mounted  skeleton  of  any  quadruped ;  the  scapula  and  ilium 
diverging,  the  humerus  and  femur  converging,  the  fore-arm  and  leg  again  diverging,  the 
foot  pointing  forward,  and  the  toes,  since  their  flexion  is  in  the  direction  opposite  to  that  of 
the  foot  at  the  ankle,  really  pointing  backward  ;  but  their  antitypes,  the  hand  and  fingers, 
seem  to  point  also  in  the  same  instead  of  the  opposite  direction,  so  that  two  and  even  three 
sets  of  muscles,  which  by  their  contraction  shorten  the  arm,  lie  upon  one  and  the  same  side 
of  the  limb. 

To  understand  this  apparent  anomaly,  it  must  first  be  remembered  that  the  entire  limbs 
are  teleological,  and  that  the  influence  of  morphology  diminishes  as  we  recede  from  the 
morphological  centre  toward  the  distal  extremities ;  moreover,  the  functions  of  the  hands 
are  various  to  the  highest  degree,  and  they,  as  the  special  agents  of  the  brain,  may  be  sup- 
posed to  pa*rtake  somewhat  of  its  independence  of  morphological  restraints. 

Embryology  throws  light  on  this  point.  In  the  early  foetal  periods,  the  two  bones  of  the 
fore-arm  are  parallel,  and  this  alone  is  sufficient  indication  of  their  morphological  relation ; 
in  the  course  of  development  the  hand  is  gradually  pronated,  so  that  the  lower  end  of  the 
radius,  the  outer  bone,  crosses  the  ulna,  and  so  becomes  internal,  causing  the  palm  to  face 
downward  and  backward,  instead  of,  downward  and  forward.  In  this  relation,  more  or  less 
firmly  connected,  the  two  bones  remain  in  quadrupeds ;  for  they,  not  being  stationary  geo- 
metrical figures,  but  organized  living  creatures  intended  to  move  from  place  to  place,  must 
be  able  to  strike  the  earth  alike  with  both  pairs  of  limbs  in  order  to  propel  the  body  in  the 
opposite  direction ;  but  in  monkeys  and  in  man,  where  the  anterior  extremities  are  not 
merely  for  progression  but  for  executing  the  higher  mandates  of  the  will,  the  radio-ulnar 
articulations  remain  free,  and  the  parts  may  be  restored  to  their  normal  condition  by  supi- 
nating  the  fore-arm,  the  palm  still  facing  downward,  but  now  also  forward  instead  of  back- 
ward ;  the  fingers  flex  forward  and  the  toes  backward,  although  made  to  act  as  continua- 
tions of  the  larger  segments,  hand  and  foot. 

Morphologically,  the  flexion  of  the  hand  at  the  wrist  must  be  in  a  direction  opposite  to 
the  flexion  of  the  fore-arm  at  the  elbow,  and  therefore  the  muscles  which  raise  the  so-called 
back  of  the  hand  toward  the  back  of  the  fore-arm  are  really  the  flexors  of  the  former  seg- 
ment, and  correspond  to  the  muscles  which  elevate  the  dorsum  of  the  foot  toward  the 
front  of  the  leg ;  and,  per  contra,  the  muscles  which  bend  the  palm  of  the  hand  toward 
the  inside  of  the  fore-arm,  are  really  extensors,  and  correspond  with  those  which  in  the 
posterior  limb  act  upon  the  foot  through  the  tendo  Achillis ;  that  is,  the  muscles  now 
called  extensores  carpi  radialis  and  ulnaris,  are  morphologically  flexors,  and  their  antagonists, 
now  called  flexores  carpi  radialis  and  ulnaris,  are  extensors,  and  will  be  so  designated  in  this 
paper. 

There  are  two  apparent  objections  to  the  above  interpretation  of  the  antitypical  rela- 


IN  THE   LIMBS  OF   MAMMALIA.  21 

tions  between  the  fore  and  hind  limbs,  but  as  will  be  seen,  they  are  only  apparent,  and 
against  their  entertainment  may  be  urged  all  that  has  been,  and  much  more  that  might  be, 
said  concerning  the  distinctions  between  morphology  and  teleology,  and  the  fallacy  of 
deductions  from  either  respecting  the  other. 

The  first  is,  that  by  rotating  back  so  as  to  leave  the  ulna  and  the  radius  parallel, 
the  former  must  correspond  to  the  inner  bone  of  the  leg,  tibia.,  and  the  latter  to  the  outer 
bone,  fibula ;  yet  the  radius  forms  most  of  the  wrist  joint,  and  the  tibia  forms  the  ankle,  and, 
as  the  power  of  rotation  gradually  disappears  in  the  mammalian  series,  it  is  the  ulna  in  the 
fore  leg,  and  the  fibula  in  the  hind  leg,  which  decreases  in  size  through  the  carnivora,  till  in 
the  permanently  pronated  fore-arm  of  the  ruminants,  solipeds,  and  some  pachyderms,  the 
ulna  is  represented  only  by  the  olecranon  process  and  the  upper  half  of  the  shaft  soldered 
to  the  radius,  while  in  the  posterior  limb  scarce  a  trace  of  the  fibula  remains ;  in  other 
words,  the  homologous  bones  of  the  two  extremities  are  developed  in  an  inverse  instead  of 
a  direct  ratio  ;  but  relations  of  more  and  less,  like  those  of  form,  are  always  dependent  upon 
function,  and  therefore  not  safe  guides  to  homology,  so  that  this  cannot  be  taken  as  a  real 
objection  to  the  law  of  longitudinality,  borne  out  as  it  is  by  the  entire  vertebrate  structure 
wherever  the  original  plan  is  retained  or  can  be  detected  under  its  various  teleological 
modifications.  We  may  however  connect  this  fact  with  another,  so  that  it  shall  to  us  seem 
to  have,  what  with  all  things  in  nature  it  certainly  must  have,  a  more  or  less  remote  foun- 
dation in  use.  As  we  have  seen  above,  in  a  morphological  point  of  view,  the  ulna  is  the 
inner,  and  the  radius  the  outer,  bone.  This  is  the  relation  which  they  bear  in  the  fcetus 
and  which  harmonizes  best  with  other  parts ;  but  this  admits  of  such  variation  that  in  nearly 
all  mammals  below  the  quadrumana,  the  radius  becomes  inner  and  the  ulna  outer.  Now, 
for  some  cause  not  yet  understood,  it  is  best  for  both  hand  and  foot  in  quadrupeds  to  be 
connected  with  the  inner  bone  of  the  legs,  and  in  provision  for  this  the  hand  is  in  all  the 
mammalia  supported  by  the  radius  which  in  the  four-footed  members  of  the  class  is  the 
inner  bone. 

The  second  and  more  obvious  though  equally  fallacious  objection  is,  that  of  the  five 
digits  which  terminate  the  anterior  and  posterior  extremities,  the  outer  ones  will  then  be 
the  thumb  and  the  little  toe,  and  the  inner  ones  the  little  finger  and  great  toe.  Perhaps 
no  correspondence  has  been  more  generally  admitted,  and  even  taken  as  a  basis  for  other 
investigations,  than  the  analogy  between  the  thumb  and  the  great  toe,  not  only  because 
in  the  commonly  accepted  condition  of  the  parts  they  both  occur  on  the  inner  sides  of  the 
hand  and  foot,  but  because  they  are  so  constantly  composed  of  two  phalanges,  while  all  the 
other  digits  possess  three. 

But  even  if  it  were  true,  which  it  is  not,  that  this  same  numerical  relation  prevailed 
among  the  three  other  classes  of  the  vertebrate  sub-kingdom,  and  there  were  therefore  some 
grounds  for  regarding  as  morphological  a  rule  for  which  no  sufficient  teleological  cause  is 
yet  apparent,  we  could  not  reasonably  accord  to  any  deviation  at  the  very  distal  extremi- 
ties of  the  limbs,  a  value  sufficient  to  outweigh  the  teachings  of  all  parts  between  them  and 
the  morphological  centre  of  the  body;  and  it  is  better  to  acknowledge  our  ignorance  of  the 
meaning  of  one  fact  than  to  purposely  ignore  the  existence  of  others  far  more  numerous 
and  important. 

I  will  here  mention  the  comparisons  between  the  anterior  and  posterior  extremities  of 
vertebrates  which,  since  the  time  of  Winslow,  (1775,)  have  been  made  by  many  of  the  most 

6 


22  WILDER  ON  MORPHOLOGY  AND  TELEOLOGY 

celebrated  anatomists,  such  as  Vicq  d'Azyr,  Soemmering,  Goethe,  Meckel,  deBlainville, 
Barclay,  Gerdy,  Blandin,  Bourgery  and  Cruveilhier,  Turenne,  Flourens,  Owen,  and  more 
recently,  Mons.  Chas.  Martins,  Professor  of  Medical  Natural  History  of  the  Faculty  of  Medi- 
cine of  Montpellier,  in  a  paper  entitled,  "  Nouvelle  comparaison  des  membres  pelviens  et 
thoraciques  chez  rhomme  et  chez  les  mammiferes  deduite  de  la  torsion  de  1'humerus," 
(Annales  des  Sciences  Naturelles,  tome  viii.  p.  45,  1857.)  And  again  in  1862,  in  a  second 
paper  entitled,  "  Memoire  sur  I'oste'ologie  compare'e  des  articulations  du  coude  et  du  genou 
chez  les  mammiferes  les  oiseaux  et  les  reptiles." 

In  the  former,  after  discussing  and  objecting  to  the  views  of  the  other  anatomists  above 
named,  he  says  on  page  55:  — 

"To  recapitulate,  these  comparisons  (paralleles)  of  the  superior  and  inferior  extremities 
of  vertebrates  may  be  reduced  to  three : 

1st.  The  hypothesis  of  Vicq  d'Azyr,  who  compares  the  superior  member  of  one  side  with 
the  inferior  member  of  the  other  side.  (Plate  ii,  figs.  1  and  3.) 

2d.  The  detailed  (detaille)  comparison  of  Bourgery,  who  combines  the  hypothesis  of 
Vicq  d'Azyr,  with  a  crossing,  in  virtue  of  which  the  head  of  the  tibia  represents  the  ulna,  its 
lower  extremity  the  radius,  while  the  femoral  extremity  of  the  fibula  corresponds  with  the 
radius,  and  its  tarsal  extremity  with  the  ulna. 

3d.  The  explanation  of  Flourens,  where  the  pelvic  member  of  one  side  is  assimilated 
with  the  thoracic  member  of  the  same  side,  the  fore-arm  being  in  a  state  of  pronation" 

We  may  easily  see,  as  Mons.  Martins  has  shown,  that  each  of  these  comparisons  is  open  to 
serious  objections,  while  their  discordance  is  such  that  even  at  this  late  day  those  anatomists 
who  do  not  utterly  discredit  the  existence  of  any  natural  relation  between  the  fore  and 
hind  limbs  "  are  in  doubt  between  them,  without  being  able  to  agree  upon  the  most  impor- 
tant point,  namely,  the  identification  of  the  two  bones  of  the  fore-arm  with  those  of 

the  leg." 

It  will  be  noted  that  through  all  these  comparisons  runs  the  effort  to  demonstrate  a 
parallelism  between  the  anterior  and  posterior  extremities,  and  not  one  of  the  anatomists, 
who  advocate  them  seems  to  have  appreciated  the  significance  of  Oken's  a  priori  assertion 
of  an  oppositeness  or  symmetry  between  the  two  ends  of  the  vertebrate  body,  which  gen- 
eralization he  simply  did  not  extend  to  the  limbs,  the  diverging  appendages  of  these  two 
regions. 

That  this  oppositeness  or  symmetry  does  really  exist,  has  I  hope  been  already  shown  in  this 
paper,  and  I  desire  to  repeat  here  that  the  first  suggestion  of  the  idea  to  me  came  from  my 
illustrious  instructor  in  anatomy,  the  Hersey  Professor  of  Anatomy  in  Harvard  University, 
from  whom,  much  rather  than  from  myself,  would  I  prefer  that  others  should  learn  what 
has  afforded  me  so  much  mental  pleasure  and  profit.1 

Mons.  Martins'  view  is  in  his  own  words,  as  follows :  "  The  humerus  is  a  bone 
twisted  on  its  axis  180  degrees.  The  femur  is  straight  without  twisting.  The  humerus 
being  a  twisted  femur,  if  we  would  compare  the  two  bones,  we  must  first  untwist  the 
humerus." 

,  In  other  words,  having  made  up  his  mind  that  the  limbs  are  parallel,  and  finding  that 
they  are  not  parallel,  he  makes  them  so  by  a  process  of  untwisting,  to  which  I  hope,  others 
will  perceive  obstacles  both  mental  and  physical. 

1  See  a  short  communication  by  Professor  J.  Wyman  to  the     ceedings  of  B.  S.  N.  H.,  vol.  vii.,  p.  317,  "  on  anterior  and 
Boston  Society  of  Natural  History  for  June  6th,  1860.     Pro-    posterior  symmetry  in  the  limbs  of  mammals." 


IN  THE  LIMBS   OF  MAMMALIA.  23 

He  then  goes  on  to  say :  "  The  anterior  extremity  having  been  thus  untwisted,  the 
radius  and  tibia  (which  he  considers  analogous  bones),  are  on  the  inside,  while  the  ulna 
and  fibula  are  on  the  outside  ;  also  the  thumb  and  great  toe,  and  the  little  finger  and  little 
toe,  are  respectively  inside  and  outside  ;"  all  which  correspondences  do  of  course  confirm 
him  in  his  preconceived  idea;  but  since,  as  has  been  shown,  they  are  only  analogies,  not 
homologies,  they  do  not  in  the  least  affect  the  true  view  of  the  case.  Anatomy  should  be 
studied  from  the  centre  outward,  as  well  as  from  the  circumference  inward. 

Under  the  head  of  "  Evidence  of  the  torsion  of  the  humerus,"  Mons.  Martins  adduces  the 
raised  line  which  passes  from  the  condyle  across  the  front  of  the  bone,  and  the  general 
direction  of  the  vessels  and  nerves  of  the  upper  arm;  not  perceiving  that  both  are  only 
physiological  provisions,  in  the  one  case  for  the  attachment  of  muscles,  and  not  appearing 
till  needed ;  and  in  the  other  for  the  better  protection  of  the  vessels  and  nerves,  as  is, 
moreover,  equally  the  case  in  the  lower  limb.  What  can  Mons.  Martins  think  of  the 
numerous  ridges  and  apparent  contortions  presented  by  all  the  bones  of  the  great  ant- 
eater  (Myrmecophaga  jubata)  ?  It  would  be  difficult  to  determine  how  many  degrees 
they  should  be  untwisted  to  conform  to  his  ideas  of  their  normal  condition.  Unfortunately, 
Mons.  Martins  has  not  taken  warning  from  his  predecessors,  and  is,  like  them,  haunted  with 
the  idea  of  parallelism,  but  his  view  has,  in  one  respect  at  least,  the  merit  of  originality ; 
for  while  they  humbly  took  things  as  they  found  them,  and  patched  them  up  as  well  as 
they  could,  he  boldly  declares  that  things  are  not  what  they  seem,  or  at  least  seem  not 
what  they  ought  to  be,  and  with  his  own  hands  sets  them  aright.  His  preestablished 
theory  is  a  very  Procrustes'  bed,  to  which  facts  must  be  adapted,  whatever  their  real 
import. 

Having  done  this  violence,  however,  his  conscience  reproaches  him,  and  he  thinks  to 
appease  outraged  Nature  by  conceding  what  he  is  pleased  to  consider  a  "metaphysical 
difficulty,"  (p.  65.)  Here,  after  admitting  that  no  such  insisting  ever  takes  place,  and  that  in 
youth,  before  the  development  of  the  muscles  has  raised  lines  on  the  bones,  no  evidence 
of  torsion  exists,  he  declares  that  "  this  torsion  of  the  humerus  is  not  mechanical,  but  only 
virtual,  though  producing  the  same  effect  as  if  it  were  real."  Also  that  "natural  history 
is  full  of  such  facts."  The  examples  he  adduces  are  undoubtedly  such ;  but  a  sounder 
illustration  is  to  be  drawn  from  the  position  of  the  whole  anterior  extremity ;  this  is  now 
admitted  by  all  to  represent  the  pair  of  ribs  with  their  diverging  appendages  of  the 
occipital  cranial  vertebra,  not  only  in  fishes  where  the  contact  is  actual,  but  also  in  all 
other  vertebrates,  man  included,  as  stated  above.  But  the  reasons  for  this  conclusion 
are  very  weighty,  since  in  addition  to  the  fact  that  the  homologous  parts  in  fishes  are 
actually  so  attached,  there  else  would  be  a  vertebral  segment  minus  a  pair  of  ribs,  and  a 
pair  of  ribs  minus  a  vertebra ;  nor  indeed  is  it  certain  that  even  in  the  higher  vertebrates 
the  anterior  extremity  is  not  at  some  time  in  contact  with  the  cranium,  while  the  other 
evidence  is  absolutely  conclusive  as  to  the  morphological  relationship.  So  that  we  may 
say  there  is  a  positive  necessity  for  so  disposing  of  these  otherwise  vagrant  vertebral  ele- 
ments. But  no  such  necessity  compels  us  to  assume  a  torsion  of  the  humerus  in  order  to 
gain  a  clear  and  philosophical  view  of  the  vertebrate  limbs ;  and  even  if  in  our  opinion  it . 
did,  no  such  torsion  ever  takes  place,  as  is  acknowledged  by  Mons.  Martins  ;  so  instead 
of  a  metaphysical  difficulty  in  the  way  of  his  supposed  reasonable  theory,  we  find 
serious,  aye,  insurmountable,  difficulties  opposed  to  a  very  metaphysical  and  visionary 
idea. 


24 


WILDER  ON  MORPHOLOGY  AND  TELEOLOGY 


Two  wrono-s  do  not  make  a  right,  and  when  we  are  in  doubt,  it  is  better  to  follow  Nature 
strictly  and  try  to  discover  her  way  of  reconciling  apparent  discrepancies,  than  to  assume 
what  is  not  the  case  in  order  to  conform  to  a  preestablished  theory. 

The  mechanical  conditions  necessary  for  making  use  of  muscular  contractility  are  two 
points,  one  generally  fixed  and  the  other  movable,  to  which  the  extremities  of  the  muscle 
are  attached,  as  in  the  face,  where  the  muscles  may  arise  from  the  bone  and  be  inserted 
directly  into  the  skin,  or  other  part  to  be  moved.  In  this  way,  of  course,  all  the  power  of 
the  muscle  is  utilized ;  but  for  the  various  requirements  as  to  rapidity  of  action  and  beauty 
of  proportion,  a  part  of  the  power  is  more  often  sacrificed  by  the  introduction  of  two  other 
points, —  one  fixed,  the  fulcrum,  and  one  movable,  like  the  part  to  be  moved,  but  nearer 
to  or  farther  from  the  fulcrum,  and  into  which  now  the  muscle  is  inserted. 

This  is  the  more  usual  method  of  applying  muscular  contraction,  and  converts  the  vari- 
ous segments  of  a  limb  into  levers,  which  are  so  far  strictly  mechanical.  Thus,  in  flexion 
of  the  fore-arm,  the  biceps,  and  brachialis  anticus  do  not  reach  directly  from  their  origins  on 
the  scapula  and  humerus  to  the  hand,  which  is  the  part  to  be  moved,  but  two  other  points 
are  introduced,  —  one  fixed  as  a  fulcrum,  the  elbow-joint,  the  other  movable  at  the  insertion 
of  the  muscles  on  the  fore-arm,  thus  between  the  fulcrum  and  the  part  to  be  moved,  which 
represents  the  weight. 

Levers  are  of  three  kinds.  The  first  is  where  the  fulcrum  lies  between  the  power  and 
the  weight ;  of  this  kind  are  all  segments  of  the  limbs  when  acted  upon  by  extensor  mus- 
cles. The  third  kind  is  where  the  power  is  applied  between  the  fulcrum  and  the  weight, 
and  to  this  class  belong  the  same  segments  when  acted  upon  by  flexor  muscles.  The 
second  kind  of  lever  is  where  the  weight  lies  between  the  power  and  the  fulcrum ;  of  this 
there  are  no  examples  in  the  body  acting  by  and  upon  itself,  but  whenever  extension  of 
a  limb  is  employed  for  raising  or  supporting  the  body  from  the  earth,  then  the  different 
segments  from  levers  of  the  first  become  levers  of  the  second  kind.  Thus,  in  rising  on 
tiptoe,  the  weight  of  the  body  rests  upon  the  ankle,  and  so  between  the  ball  of  the 
foot,  which,  resting  on  the  earth,  forms  the  fulcrum,  and  the  heel,  to  which  by  the 
tendo  Achillis  is  applied  the  power  of  the  gastrocnemius  and  soleus  muscles.  I  subjoin  line 
illustrations  of  these  three  kinds  of  levers. 


Adduction  and  abduction  are  only  lateral  flexion  and  extension;  circumduction  is  a  com- 
pound movement  resulting  from  gradual  and  successive  direct  and  lateral  flexion  and 
extension ;  even  rotation  is  essentially  a  peculiar  form  of  the  same  two  movements,  for  the 
power  is  applied  at  the  periphery  of  the  rotated  bone  or  limb,  and  so  at  one  end  of  an  imag- 
inary diameter  line  through  the  centre,  which  latter  represents  the  fulcrum ;  the  weight 
may  be  regarded  either  as  suspended  from  the  half  of  the  line  or  lever  furthest  from  the 
power,  in  which  case  the  diameter  line  would  be  a  lever  of  the  first  kind,  or  as  sustained 


IN  THE   LIMBS   OF   MAMMALIA.  25 

upon  the  nearer  half  and  thus  between  the  power  and  the  fulcrum,  when  it  would  be  a 
lever  of  the  second  kind. 


Of  course,  any  obliquity  in  the  direction  of  the  muscle  or  bone,  or  any  variation  in  the 
shape  of  the  rotated  bone  or  limb,  or  the  presence  of  any  part  as  the  hand,  which  may  be 
appended  to  the  extremity,  will  only  appear  to  complicate  the  motion  without  essentially 
affecting  its  character. 

The  general  law,  according  to  which  the  muscles  of  the  mammalian  limb  seem  to  be 
arranged,  is  in  no  way  startling  or  peculiar,  and  the  only  wonder  is  that  it  has  never  been 
observed  before.  No  new  facts  are  required  for  its  illustration ;  human  anatomy  contains 
the  great  bulk  of  the  material,  though  of  course  aided  by  comparative  anatomy;  and  it 
needs  only  to  see  that  the  facts  are  not  arbitrary,  but  bear  such  mutual  relation  as  clearly 
to  admit  of  one  general  statement.  I  will  commence  with  the  muscles  which  act  upon  the 
fore-arm,  that  segment  of  the  anterior  extremity  in  which  I  first  detected  the  arrangement, 
and  in  which  it  is  quite  closely  adhered  to. 

There  are  two  direct  flexor  muscles  of  the  fore-arm  :  one  is  the  biceps  arising  from 
the  scapula  and  inserted  into  the  radius,  the  other  is  the  brachialis  anticus  arising  from  the 
humerus  and  inserted  into  the  ulna,  though  both,  of  course,  are  attached  to  one  and  the 
same  segment,  fore-arm.  But  this  segment  may  be  also  flexed  indirectly  by  the  ulnar  and 
radial  extensors  of  the  wrist ;  for  when  a  joint  is  partially  or  wholly  fixed  by  the  simul- 
taneous contraction  of  antagonist  muscles  on  opposite  sides,  then  the  two  segments  between 
which  the  joint  intervenes  become  teleologically  as  one  segment,  which  is  acted  upon  by  all 
muscles  arising  from  any  other  segment,  and  inserted  into  either  of  them;  each  segment 
being  acted  upon  directly  by  the  muscles  which  are  attached  to  itself,  and  indirectly  by  those 
attached  to  the  other.  The  fore-arm  is  directly  extended  by  two  muscles,  one  short,  the 
humeral  heads  of  the  triceps,  and  the  other  long,  the  scapular  head  of  the  same  ;  the  indirect 
extensors  are  the  ulnar  and  radial  flexors  of  the  wrist,  commonly  called  extensores  carpi  ul- 
naris  and  radialis. 

The  humerus  is  flexed  upon  the  scapula  by  three  muscles,  —  directly  by  two,  one  short, 
teres  major,  arising  from  the  scapula,  and  one  long,  arising  upon  the  side  of  the  body, 
latissimus  dorsi ;  and  indirectly  by  the  third,  the  scapular  head  of  the  triceps  which  is  also  the 
long  direct  extensor  of  the  segment  next  below. 

How  is  it  now  with  the  corresponding  segments  of  the  posterior  extremity  ?  The  leg  is 
flexed  at  the  knee,  directly  by  two  sets  of  muscles,  one  of  which  is  represented  by  the  popli- 
tceiis  with  the  short  head  of  the  biceps  when  it  exists,  the  former  corresponding  with  the 
short  direct  flexor  of  the  fore-arm,  brachialis  antkus ;  the  other  set  is  composed  of  the  long 
head  of  the  biceps  with  its  accessories,  the  semi-tendinosus  and  semi-membranosus,  the  former 
corresponding  with  the  long  direct  flexor  of  the  fore-arm,  biceps  himicri  ;  while  the  peronceus 
long  us  and  tibialis  posticus,  with  their  accessories;  the  muscles  composing  the  calf  of  the  leg, 
and  which  are  long  direct  extensors  of  the  foot,  are,  if  the  ankle-joint  is  fixed,  the  indirect 
flexors  of  the  leg,  the  segment  next  above,  and  correspond  thus  with  the  extensores  carpi 
and  the  palmaris  in  the  other  extremity. 

7 


26  WILDER  ON  MORPHOLOGY  AND  TELEOLOGY 

Here  are  enough  examples  to  illustrate  the  general  rule  which  may  be  stated  in  several 
ways,  according  as  we  specially  regard  the  muscles  themselves,  or  the  bones ;  and  these  as 
segments  of  insertion  acted  upon,  or  as  segments  of  origin  from  which  the  muscles  act. 

The  muscles  of  the  limbs  form  two  groups,  long  and  short ;  the  short  muscles  arising  from 
theirs/  segment  above  that  into  which  they  are  inserted,  and  the  long  from  the  second 
segment  above.  A  short  muscle  can  act  in  but  one  way,  but  a  long  one  has  two  actions  : 
one  direct  upon  the  segment  into  which  it  is  inserted,  the  other  indirect  upon  the  segment 
which  intervenes  between  its  origin  and  insertion. 

Each  segment  of  a  limb  is  flexed  or  extended  by  three  muscles  or  groups  of  muscles;  of 
these,  two  are  inserted  into  itself,  and  are  therefore  called  direct  muscles,  flexors  or  extensors  ; 
one  arising  from  the  segment  next  above  the  segment  to  be  moved,  and  hence  called  the 
short  direct,  and  the  other  arising  from  the  second  segment  above,  and  hence  called  the  long 
direct ;  the  third  muscle  is  termed  the  indirect  flexor  or  extensor,  and  is  the  long  direct  exten- 
sor or  flexor  respectively  of  the  next  segment  below,  and  between  whose  origin  and  inser- 
tion intervenes  the  segment  under  consideration. 

Or  it  may  be  stated  in  yet  a  third  way,  using  the  humerus  as  an  example.  Each  seg- 
ment not  only  gives  insertion  to  the  four  muscles  which  act  upon  itself,  short  and  long 
extensors,  deltoid  and  pectoralis  major,  and  flexors,  teres  major  and  latissimus  dorsi,  but 
also  affords  origin  to  four  more,  of  which  two  are  the  short  flexor  and  extensor  of  the  seg- 
ment next  below,  brachialis  anticus  and  humeral  heads  of  triceps,  and  two,  the  long  extensor 
and  flexor  of  the  second  segment  below,  extensor  and  flexor  carpi  radialis,  which  are  also  the 
indirect  extensor  and  flexor,  respectively,  of  the  first  segment  below. 

There  are  several  advantages  apparent  in  this  arrangement  of  the  muscles  of  the  limbs, 
and  probably  many  more  than  have  yet  occurred  to  me.  The  most  comprehensive  is,  that 
while  one  half  of  the  muscles,  the  short,  are  able  to  execute  but  one  movement,  since  but 
one  joint  intervenes  between  their  origin  and  insertion,  the  other  half  may  act  in  either  of 
two  ways,  since  two  joints  intervene  between  their  origin  and  their  insertion.  Take  for 
example  the  biceps  humeri ;  its  lower  extremity  is  associated  with  the  short  flexor  of  the 
fore-arm,  brachialis  anticus,  which  is  thus  its  associate  of  insertion,  while  its  upper  extremity  is 
associated  with  the  short  extensor  of  the  humerus,  deltoid,  which  is  thus  its  associate  of 
origin ;  in  like  manner  its  antagonist  of  origin  is  the  short  flexor  of  the  humerus,  teres  major, 
and  its  antagonist  of  insertion,  the  short  extensor  of  the  fore-arm,  humeral  heads  of  triceps. 
Now  when  the  elbow,  its  joint  of  insertion,  is  fixed  by  the  simultaneous  contraction  of  its 
associate  of  insertion  and  its  short  antagonist  of  insertion,  then  the  biceps  can  act  only  with 
the  deltoid,  its  associate  of  origin  ;  but  when  the  shoulder,  its  joint  of  origin,  is  fixed  by  the 
simultaneous  contraction  of  its  associate  of  origin  and  short  antagonist  of  origin,  then  the 
biceps  acts  with  its  associate  of  insertion  to  flex  the  fore-arm. 

Here  is  the  great  advantage  in  having  both  long  and  short  muscles ;  each  long  one  may 
act  in  two  ways,  and  the  short  ones  are  then  required  to  counteract  one  of  the  actions 
while  the  other  is  performed.  Moreover,  it  often,  and  in  fact  usually,  happens,  that  two  or 
more  segments  of  a  limb  are  to  be  flexed  or  extended  together,  and  this  is  provided  for  by 
the  same  arrangement,  with  the  additional  fact  that  the  short  muscles  are  generally  shorter 
and  thicker  than  the  long.  Suppose  that  two  contiguous  segments,  as  arm  and  fore-arm, 
are  to  be  extended  at  the  same  time.  In  this  case  neither  of  the  short  flexors,  teres  major 
and  brachialis  anticus,  can  act  at  all,  but  the  long  flexor  of  the  lower  segment,  biceps,  is  ena- 
bled to  act  as  the  indirect  extensor  of  the  upper  segment  with  its  short  associate  of  origin, 
deltoid,  its  flexor  power  being  counteracted  by  its  short  antagonist  of  insertion,  humeral 


IN   THE   LIMBS   OF   MAMMALIA.  27 

heads  of  triceps,  which,  being  more  powerful,  continues  to  act  as  an  extensor,  beside  leaving 
the  long  extensor,  scapular  head  of  triceps  entirely  free  ;  the  action  of  the  latter,  as  an  indirect 
flexor  of  the  humerus,  being  in  like  manner  more  than  counteracted  by  the  short  extensor 
of  the  same,  deltoid.  But  if  the  same  two  segments  are  to  be  simultaneously  flexed,  then 
the  short  extensors  do  not  act  at  all,  while  the  extensor  power  of  the  long  direct  flexor, 
biceps,  is  overbalanced  by  its  short  antagonist  of  origin,  teres  major,  and  that  of  the  long 
direct  extensor,  scapular  head  of  triceps,  by  its  short  antagonist  of  insertion,  brachialis  anticus, 
leaving  the  two  long  flexors,  latissimus  dorsi  and  biceps,  free  as  flexors  of  the  two  segments ; 
moreover,  the  long  rotators  of  the  fore-arm,  supinator  longus  and  pronator  teres,  with  the  long 
extensor  of  the  wrist,  extensor  carpi  radialis,  and  here  by  exception,  even  the  long  flexor  of 
the  third  segment  below,  flexor  communis  digitorum  sublimis,  now  act  as  indirect  flexors  of  the 
fore-arm,  their  direct  actions  being,  if  necessary,  prevented  by  their  short  antagonists,  or  in 
the  case  of  the  rotators  by  each  other. 

Again,  as  has  perhaps  been  already  inferred,  the  three  muscles  acting  upon  any  one  seg- 
ment, are,  potentially  or  morphologically,  of  the  three  degrees  of  length  in  direct  ratio,  but  of 
thickness  in  inverse  ratio  ;  and  since  the  power  of  a  muscle  is  in  proportion  to  its  thickness, 
and  the  distance  through  which  it  can  contract,  according  to  the  length  of  its  fibres,  it  fol- 
lows that  a  movement  is  most  rapid  and  forcible  at  the  beginning,  when  all  three  muscles 
act  together,  and  least  so  at  its  close  when  the  longer  and  weaker  muscles  act  alone ;  and 
this  agrees  with  the  observations  on  the  variation  in  the  force  of  contraction  of  a  single 
muscle,  which  is  found  to  be  most  forcible  at  the  beginning,  least  so  at  the  end. 

I  have  now  stated  the  general  law  and  the  general  advantages  gained  thereby  as  illus- 
trated in  those  regions  where  the  muscles  are  most  familiar  and  where  the  law  is  quite 
closely  adhered  to,  yet  even  here  were  some  departures  from  it ;  and  in  the  remaining  por- 
tions of  the  limbs  will  be  found  even  more  variations,  according  to  the  number  and  kind  of 
movements  required  at  the  several  joints. 

There  is  considerable  doubt  with  regard  to  the  muscles  acting  upon  the  scapula,  and  for 
two  principal  reasons,  one  teleological  and  the  other  morphological ;  for,  beside  the  compli- 
cation necessary  for  the  very  free  movement  upon  the  walls  of  the  thorax,  of  a  part  which 
in  many  mammalia  has  no  bony  connection  therewith,  there  is  so  much  difficulty  in  compre- 
hending the  true  relations  of  some  muscles  arising  from  the  head  and  cervical  vertebrae, 
and  inserted  into  the  scapula  and  clavicle,  as  to  afford  additional,  though  for  the  present 
vague,  evidence  concerning  that  remarkable  change,  by  which  the  scapular  arch  has  been 
displaced  backward  from  the  occipital  cranial  vertebra,  of  which  it  is  in  most  fishes  actu- 
ally, and  in  all  vertebrates  morphologically,  the  pair  of  ribs. 

The  scapula  slides  upon  the  sides  of  the  thorax,  and  is  separated  from  it  chiefly  by  mus- 
cles, having  in  most  mammalia  no  bony  connection  with  the  rest  of  the  skeleton  ;  and  the 
muscles  which  act  directly  upon  it,  are,  like  those  of  the  face,  inserted  at  once  into  the  part 
to  be  moved,  without  the  additional  mechanical  complication  of  levers.  It  may  be  elevated 
and  depressed,  drawn  forward  toward  the  sternum,  or  backward  toward  the  vertebral 
column.  Two  direct  muscles  draw  the  scapula  forward;  one,  the  pectoralis  minor,  inserted 
into  the  coracoid  process,  the  other,  the  serratus  magnus,  attached  along  the  posterior  bor- 
der, both  arising  from  the  ribs ;  two  direct  muscles  also  draw  it  backward,  the  trapezius 
outside,  inserted  upon  the  clavicle  and  spine  of  the  scapula,  and  the  rhomboideus  inside,  in- 
serted into  the  posterior  border,  and  both  these  arising  from  the  vertebral  column.  Here 
then,  as  usual,  are  two  flexors  and  two  extensors,  but  as  there  is  no  part  beyond  the  verte- 


28 

bral  column  or  sternum  on  the  middle  line,  all  four  are  ideologically  short  muscles,  since  but 
one  joint  intervenes  between  their  origins  and  insertions  ;  it  seems  likely,  however,  that  in  a 
morphological  point  of  view,  the  outer  ones,  trapezius  and  pedoralis  minor,  are  the  long  mus- 
cles. The  scapula  is  elevated  and  depressed  directly  by  the  oblique  descending  and  ascend- 
ing fibres  of  the  muscles  already  described  or  indirectly  by  the  long  muscles  of  the  humerus 
pedoralis  major  and  latissimus  dor  si.  The  true  affinities  of  the  cleido-mastoid  and  levator  clavic- 
ulce  have  not  yet  been  determined  ;  the  omo-ht/oid  is  probably  the  representative  of  an 
intercostal  muscle  connecting  the  ribs  of  the  occipital  and  parietal  vertebras  ;  the  leva- 
tor  anguli  scapidoe  is  commonly  regarded  as  a  distinct  muscle,  but  from  some  facts  in  its 
comparative  anatomy,  it  appears  to  me  rather  as  a  dismemberment  of  the  serratus  magnus. 
All  this  is  rather  unsatisfactory,  and  will,  I  think,  continue  so  till  embryology  and  com- 
parative anatomy  have  demonstrated  the  true  relations  of  these  muscles,  and  the  extent  to 
which  they  vary  from  the  common  type  of  the  intercostales  ;  for  it  must  be  remembered, 
that  the  scapula  and  its  posterior  representative,  the  ilium,  are  not,  properly  speaking,  seg- 
ments of  the  limbs,  but  the  dorsal  moities  or  pleurapophyses  of  modified  ribs,  at  the  junction 
of  which  with  the  ventral  moities  or  hcemapophyses,  are  given  off  diverging  appendages  in 
the  shape  of  the  anterior  and  posterior  extremities. 

The  deltoid  is  the  short  extensor  of  the  humerus,  arising,  as  it  should,  from  the  scapula ; 
either  the  long  extensor  is  wanting,  or,  as  is  more  probable,  it  is  represented  by  the  pedo- 
ralis major,  or  one  of  the  several  subdivisions  of  the  latter,  existing  in  quadrupeds,  where 
the  clavicle  is  deficient.  The  long  and  short  flexors  of  the  arm,  latissimus  dorsi  and  teres 
major,  have  been  already  noticed.  With  regard  to  the  supra  and  infra  spinati,  the  subscapu- 
laris  and  teres  minor,  I  am  not  yet  decided ;  in  man  they  are  chiefly  rotators  of  the  limb  at 
the  shoulder,  but  in  quadrupeds  where  the  humerus  is  simply  flexed  or  extended,  the 
three  former  have  more  or  less  power  of  extension,  while  the  teres  minor  is  a  flexor.  The 
muscles  acting  upon  the  fore-arm  have  been  already  described ;  the  dorso-epitrochlien  of 
Duvernoy  ("  Des  caracteres  anatomiques  des  grands  singes  pseudo-anthropomorphes," 
Archives  du  Museum,  vol.  viii.)  which  exists  in  the  quadrumana  and  most  of  the  mamma- 
lia, has  nearly  the  same  relations  as  the  scapular  head  of  the  triceps,  being,  when  inserted 
upon  the  olecranon,  a  dire'ct  extensor  of  the  fore-arm  as  well  as  an  indirect  flexor  of  the 
humerus,  which  alone  it  flexes  directly  when  attached  to  the  internal  condyle. 

The  relations  of  the  biceps  in  the  higher  mammalia  are  somewhat  complicated  by  the  pro- 
visions for  rotation  between  the  two  bones  of  the  fore-arm,  converting  that  otherwise 
single  segment  of  the  limb  into  two,  whose  greater  lengths  are  parallel  with,  instead  of 
perpendicular  to,  the  axis  of  motion  at  the  radio-ulnar  articulation  ;  and  since  the  biceps  is 
attached  to  the  radius,  it  really  extends  over  three  joints,  and  can  act  in  three  ways :  at  the 
shoulder  in  extension  of  the  humerus,  at  the  elbow  in  flexion  of  the  fore-arm,  and  at  the 
radio-ulnar  articulation  in  supination  or  extension  of  the  radius ;  for,  regarding  these  two 
bones  alone,  supination  is  extension,  and  pronation  is  flexion. 

It  will  be  seen,  however,  that  the  radio-ulnar  articulation  is  not  a  joint,  in  the  same  sense 
as  is  the  shoulder  or  the  elbow.  These  latter  are  constant  or  nearly  so  throughout  the 
vertebrate  series,  and  are  important  anatomical  characters ;  they  are  strictly  morphological 
joints ;  but  the  arrangement  by  which  movement  is  permitted  between  the  two  bones  of 
the  single  segment  fore-arm,  exists  in  comparatively  few  species  of  mammalia,  and  these 
such  as  in  their  approach  to  man,  have  their  fundamental  structure  most  modified  in  rela- 
tion to  the  higher  uses  they  are  to  perform.  The  radio-ulnar  articulation  is  properly  a 
tekobgical  joint,  and  as  might  be  expected,  its  existence  does  not  really  interfere  with  the 


IN  THE   LIMBS  OF  MAMMALIA. 

of  the  fingers,  unless  the  proper  extensors  of  the  first,  second,  and  fifth  digits  are  remaining 
portions  of  it ;  the  only  common  extensor,  like  the  superficial  flexor,  having  been  moved 
as  to  its  origin,  two  segments  above  its  normal  position,  like  the  short  extensor. 

In  general,  the  muscles,  like  the  bones  of  the  posterior  extremities,  repeat  in  an  opposite 
direction  those  of  the  anterior,  but  the  exact  correspondences  between  individual  muscles 
are  far  more  difficult  to  determine  than  would  be  expected,  either  from  the  apparently 
more  simple  functions  of  the  limbs  as  such,  or  from  the  close  antitypy  exhibited  by  their 
osseous  framework.  Moreover,  extended  and  minute  comparisons  of  parts  have  as  yet 
been  almost  wholly  confined  to  anthropotoiny,  a  branch  of  comparative  anatomy  which 
treats  of  a  structure  teleologically  most  perfect,  but  morphologically  monstrous. 

The  legs  are  usually  regarded  as  very  simple  and  regular  in  their  structure,  because 
their  only  function  is  that  of  locomotion ;  but  it  is  from  this  very  cause  that  they  are  more 
complicated  than  the  corresponding  limbs  of  quadrupeds,  for  they  are  also  the  only  organs 
of  locomotion,  and  therefore  are  required  to  maintain  the  equilibrium  of  the  erect  body 
during  progression,  a  duty  which  in  the  four-footed  members  of  the  class,  is  shared  by  the 
anterior  extremities. 

By  a  sort  of  favoritism  shown  to  the  anterior,  and  physiologically  more  noble  region  of 
the  body,  when  the  functions  of  prehension  are  transferred  from  the  quadrupedal  head  to 
its  diverging  appendages,  the  arms,  these  are  in  their  turn  allowed  to  impose  upon  their 
posterior  representatives  their  share  in  sustaining  and  propelling  the  body  ;  but  the  latter, 
less  highly  favored,  can  merely  make  an  awkward  protest  when  the  kangaroo  throws  upon 
the  base  of  his  huge  caudal  appendage  the  task  of  supporting  his  body  while  he  kicks  out 
with  his  hind  legs,  and  when  certain  monkeys  suspend  themselves  by  the  other  end  of  their 
tails  so  as  to  leave  all  their  limbs  free. 

The  natural  position  of  the  quadruped  is  sustained  in  part  according  to  physical  laws,  as 
is  a  table  upon  its  four  pedestals ;  but  that  of  man  is  in  direct  defiance  of  these  laws,  which 
are  ever  ready  to  assert  their  rights  the  moment  his  will  ceases  to  guard  against  them ; 
and  he,  the  lord  of  creation,  is  obliged  to  take  his  rest  in  an  attitude  other  than  that  in 
which  his  superiority  is  exercised ;  his  elevation  is  great,  but  according  to  the  law  of  the 
succession  of  extremes,  his  fall  is  proportionately  great,  when  the  means  of  that  elevation 
are  withdrawn ;  he  pays  as  it  were,  a  physiological  price  for  his  physiological  preeminence. 

As  in  the  arms  of  man  were  found  extra  muscles  and  articulations,  with  direct  reference 
to  the  superadded  function  of  rotation,  so  in  the  lower  limbs  there  appears  to  be  an 
increase  in  the  number  of  muscles,  and  changes  in  the  relation  of  those  which  are  morpho- 
logically entitled  to  be  present,  in  view  of  the  fact  that  by  two  legs,  instead  of  four,  is  the 
body  to  be  propelled  and  its  equilibrium  maintained ;  therefore,  they  cannot  be  regarded 
as  presenting  the  normal  condition  of  parts  for  correcting  our  ideas  as  derived  from  the 
arms,  since  they  are  quite  as  fully,  though  less  obviously  and  directly,  under  teleological 
influence. 

The  real  extent  of  the  antitypical  relations  between  the  anterior  and  posterior  extremi- 
ties must  be  learned  from  those  vertebrates  in  whom  they  are  actually  fore  and  hind;  and 
for  this,  when  time  and  opportunity  allow,  such  works  as  that  of  Strauss-Durckheim  on 
u  the  Anatomy  of  the  Cat ; "  of  Della-Chiaje  on  that  of  the  Testudo  Europaea ;  and  of  Meckel 
on  that  of  the  Ornithoryncus,  with  the  more  comprehensive  work  of  the  latter  anatomist, 
«  Traite  generale  d' Anatomic  comparee,"  will  be  invaluable.  Prior  to  these  investigations, 
any  such  inferences  as  may  now  be  drawn  from  the  human  structure  must  be  regarded  as 
provisional  and  by  no  means  as  conclusive. 


WILDER  ON  MORPHOLOGY  AND  TELEOLOGY 

general  plan  of  the  muscles  any  more  than  if  the  fore-arm  were,  in  some  one  species, 
broken,  and  thus  movable  at  the  middle  of  its  length,  some  of  the  muscles  being  inserted 
upon  the  proximal,  and  others  upon  the  distal  moiety.  Moreover,  the  muscles  specially 
connected  with  this  joint,  lie  all  upon  the  same  side  of  the  limb  ;  namely,  with  the  flexors 
of  the  fore-arm  and  the  extensors  of  the  hand,  while,  if  it  were  a  real  joint,  we  should 
expect  to  find  the  supinators  or  extensors  on  the  same  side  as  the  flexors  of  the  fore-arm, 
and  the  pronators  or  flexors  on  the  same  side  with  the  extensors  of  the  hand ;  but  in  fact, 
as  before  stated,  they  all  lie  upon  the  same  side ;  whereby  the  long  supinator  and  pronator 
are  both  made  to  assist  the  flexion  of  the  fore-arm,  which  is  evidently  of  more  importance 
than  its  extension.  The  radio-ulnar  articulation  is  therefore  a  teleological  interpolation, 
presenting  none  of  the  characters  of  a  true  joint  between  the  real  segments  of  a  limb. 

The  two  opposite  movements  of  pronation  and  supination  are  performed  by  two  sets  of 
muscles ;  those  of  one  set  are  short,  the  pronator  quadratus  and  supinator  brevis,  arising  from 
the  ulna,  the  first  segment  above  their  insertion  ;  those  of  the  other  set  are  long,  the  pro- 
nator teres  and  supinator  longus,  arising  from  the  humerus  or  second  segment  above  their 
insertion ;  and  here,  as  with  the  muscles  of  the  true  joints,  the  short  muscles  have  but  one 
action,  to  move  the  radius  or  counteract  each  other ;  while  the  long  ones  either  act  directly 
in  a  similar  manner  or  indirectly  flex  the  fore-arm,  the  segment  which,  as  to  its  fixed  bone, 
the  ulna,  intervenes  between  their  origins  on  the  humerus  and  their  insertions  on  the 
radius.  The  biceps  is  really  the  most  powerful  supinator  of  the  radius,  and  can  turn  the 
palm  of  the  hand  completely  upward;  while  the  two  supinatores  can  only  bring  the 
thumb  or  radial  border  of  the  hand  uppermost.  There  are  four  muscles  acting  directly 
upon  the  hand,  (carpus  and  metacarpus ;)  two  flexors,  radial  and  ulnar,  and  two  exten- 
sors, radial  and  ulnar ;  the  ulnar  flexor  and  extensor  arise  chiefly  from  the  bones  of  the  fore- 
arm, and  are  the  short  muscles,  since  they  come  from  the  first  segment  above  that  to  be 
moved,  and  the  two  radial  are  the  long  muscles  arising  from  the  humerus,  the  second 
segment  above.  The  long,  or  radial  extensor,  is  also  the  indirect  flexor  of  the  fore-arm, 
and  perhaps,  morphologically,  the  radial  flexor  is  the  indirect  extensor  of  the  same  inter- 
mediate segment,  though  I  am  not  aware  that  it  ever  is  so  actually ;  and  on  the  contrary, 
when,  as  in  the  horse,  the  humeral  condyles  are  wanting,  the  extensor  arises  on  the 


inner 


side,  below  the  centre  of  motion,  and  would  thus  act  as  an  extensor  of  the  fore-arm,  while  the 
flexor  arises  from  the  outer  side  above  the  centre  of  motion,  and  is  therefore  physiologically 
a  flexor  of  the  same  segment. 

If  we  regard  the  fingers  as  a  single  segment,  then  there  is  a  short  and  a  long  flexor, 
flexor  communis  sublimis  and  flexor  communis  profimdus ;  but  these  would  then  seem  to  arise 
each  from  one  segment  above  what  would  be  expected ;  the  former  from  the  humerus, 
and  the  latter  from  the  fore-arm ;  but  the  insertion  of  a  muscle  is  of  more  importance  mor- 
phologically than  its  origin,  which  latter  is  in  relation  to  the  required  length  of  the  muscle 
and  the  proportion  of  parts,  and  therefore  more  liable  to  vary ;  so,  taking  the  less  modified 
foot  for  comparison,  it  seems  more  natural  to  suppose  that  the  flexor  profundus  is,  morpho- 
logically, the  long  flexor,  and  the  flexor  sublimis  the  short ;  its  origin  having  been  moved 
upward  two  segments  on  account  of  the  very  extensive  movements  required  of  the  fingers, 
more  extensive  in  proportion  to  their  size  than  those  of  any  other  part  of  the  body. 

With  this  view,  the  short  flexor  of  the  fingers,  like  the  short  flexor  and  extensor  of  the 
toes,  is  inserted  into  the  second  phalanges.  That  the  proper  short  palmar  muscles  of  the 
thumb  and  little  finger  are  not  the  true  representatives  of  the  short  flexor  is  shown  by 
their  co-existence  in  the  foot  with  an  unmistakable  short  flexor.  There  is  no  long  extensor 


IN  THE   LIMBS   OF   MAMMALIA.  31 

It  will  now  be  shown  to  what  extent  our  teleological  morphology,  as  it  were,  of  the  long 
and  the  short  muscles  has  been  traced  in  the  lower  limbs  of  man.  The  ilium,  unlike  its 
longitype,  the  scapula,  is  firmly  attached  to  the  vertebral  column,  and  does  not,  even  in 
appearance,  constitute  a  segment  of  the  limb.  Physiologically,  the  short  and  long  direct 
flexors  of  the  thigh  are  the  iliacus  and  the  psoas  magnus ;  but  the  former  is  without  doubt 
the  morphological  representative  of  the  subscapularis,  which  is  in  man  an  internal  rotator, 
but  in  most  other  mammalia  an  accessory  flexor  of  the  humerus ;  and  the  real  antitype  of 
the  teres  major  is  the  small  muscle  first  described  under  the  name  of  scansorius  by  Traill, 
("  Observations  on  the  Anatomy  of  the  Orang-outang  (Chimpanzee)"  Memoirs  of  the  Werne- 
rian  Natural  Historical  Society,  vol.  iii.  Feb.  7,  1818 ;)  afterward  by  Prof.  Owen,  ("Myology 
of  Simia  satyrus,"  Proceedings  of  Zoological  Society  of  London,  January  25  and  May  30, 
1831,)  and  lately  by  me,  ("  Contributions  to  Comparative  Myology  of  Chimpanzee,"  Boston 
Journal  Natural  History,  page  369,  vol.  vii.) 

The  long  flexor  of  the  humerus,  latissimns  dorsi,  seems  to  have  no  posterior  representa- 
tive ;  the  glutceus  maximus  at  once  suggests  the  deltoid,  and  may  be  the  morphological  as 
well  as  the  physiological  short  extensor  of  the  femur;  while  perhaps  that  portion  of  it  which 
arises  from  the  sacrum,  represents  the  long  extensor,  corresponding  with  some  one  of  the 
subdivisions  of  the  trapezius  of  quadrupeds  ;  for  it  must  be  remembered  that  when  in  them 
the  clavicle  is  wanting,  certain  contiguous  portions  of  the  trapezius  and  deltoid  become 
continuous. 

The  glutceus  medius  and  psoas  magnus  are  suggestive  of  the  supra  and  infra  spinati,  and  the 
glutceus  minimus  of  the  teres  minor ;  the  pectincws  represents  the  pectoralis  major,  and  the  ad- 
ductores  brevis,  longus  and  magnus,  are  only  excessive  developments  of  what  the  coraco-brachialis 
is  anteriorly  ;  their  great  size  in  man  being  in  evident  relation,  not  only  to  his  erect  position 
but  also  to  his  firm  seat  on  the  noble  animal  which  is  to  him  strength  and  speed  ;  but  in 
the  ape  the  rami  of  the  ischia  and  pubes  are  lengthened  downward,  and  so  the  adductors 
arising  therefrom  act  powerfully  as  extensors  of  the  limb  in  leaping. 

The  close  antitypy  between  the  direct  extensors  of  the  leg  and  those  of  the  fore-arm 
has  always  been  remarked,  and  Cruveilhier  even  gave  to  the  former  the  name  "  triceps 
femoralis."  The  rectus,  like  the  scapular  head  of  the  triceps  humeralis,  is  also  the  indirect 
flexor  of  the  femur,  the  segment  which  intervenes  between  its  origin  on  the  ilium  and  its 
insertion  upon  the  inner  bone  of  the  leg ;  the  patella,  a  sesainoid  bone  developed  in  the 
tendon  of  the  extensor,  is  generally  considered  to  be  the  longitype  of  the  olecranon  proc- 
ess, both  from  their  ordinary  relations  to  the  tendons  of  the  muscles  and  from  the  fact  that 
the  latter  is  developed  from  a  distinct  osseous  centre,  and  does  in  man  sometimes  exist  as 
a  separate  piece  connected  to  the  shaft  of  the  ulna  by  the  continuation  of  the  tendon. 
The  sartorius  resembles  the  muscle  called  "  epitrochlien"  by  Duvernoy,  and  already  referred 
to  in  this  paper ;  but  the  former,  since  there  is  no  long  flexor  of  the  femur,  takes  its  origin 
from  the  anterior  superior  spinous  process  of  the  ilium. 

But  if  the  direct  extensors  of  the  leg  are  morphologically  satisfactory,  the  flexors  are 
quite  the  reverse,  and  in  them  is  at  once  seen  the  effect  of  the  twofold  duties  imposed 
upon  the  lower  limbs;  the  muscles  are  too  numerous,  and  most  of  them  are  long  ones.  The 
semi-membranosus,  semi-tendinosus  and  gracilis  attached  to  the  tibia  are  apparently  all  acces- 
sory to  the  long  head  of  the  Uceps,  which,  inserted  upon  the  fibula  alone,  is  probably  the 
real  long  flexor  of  the  leg,  and  thus  the  antitype  of  the  Uceps  humeri,  as  the  short  head  of 
the  Uceps,  when  it  exists,  is  accessory  to  the  poplitceus  which  is  the  antitype  of  the  brachialis 
anticus,  and  thus  the  short  flexor  of  the  leg.  Ordinarily,  all  the  muscles  of  the  lower 


32  WILDER  ON  MORPHOLOGY  AND  TELEOLOGY 

extremity  take  their  fixed  point  of  action  at  the  distal  instead  of  the  proximal  ends  of  the 
several  segments,  and  the  accessory  long  flexors  of  the  leg  are  evidently  more  important 
as  mutually  counteracting,  and  thus  as  balancing  flexors  and  extensors  of  the  pelvis,  than  as 
either  direct  flexors  of  the  leg  or  indirect  extensors  of  the  thigh. 

The  direct  extensors  of  the  foot  are,  in  man,  very  large,  and  two  of  them,  the  gastroc- 
ncmius  and  the  soleus,  with  the  enormously  developed  tarsal  bone,  upon  which  they  are 
inserted,  seem  to  bear  relation  to  the  above-named  physiological  necessity ;  however,  they 
are  very  good  long  and  short  muscles  accessory  to  the  tibialis  posticus  and  the  peronceus 
lonyus,  which  seem  to  be  the  morphological  short  and  long  extensors  j  and  in  the  A'i, 
(Bradypus  tridactylus,)  the  latter  muscle  does  actually  take  part  of  its  origin  above  the 
knee-joint* 

The  short  flexor  of  the  foot  is  the  tibialis  anticus,  and  the  long  the  peronceus  brevis,  which 
in  man  is  also  made  to  act  as  an  extensor  by  its  tendon  passing  behind  the  outer 
malleolus,  which,  however,  like  the  lower  extremity  of  the  fibula,  is  by  no  means  constant 
in  mammalia. 

The  toes  are  provided  with  a  short  and  a  long  flexor,  — flexor  brevis  digitorum,  and  flexor 
longus  digitorum,  —  and  with  a  short  and  long  extensor  with  corresponding  names  ;  but  the 
morphology  of  these,  and  their  relations  to  those  of  the  fingers,  wrill  be  more  easily  learned 
from  some  animals  in  which  there  is  more  resemblance  between  the  carpus  and  tarsus 
without  any  such  prominence  of  one  bone  of  the  latter,  as  in  most  mammalia,  and  espe- 
cially in  man. 

The  following  table  may  serve  as  a  contribution. toward  a  more  complete  understanding 
of  the  correspondences  between  the  muscles  of  the  anterior  and  posterior  extremities:-— 


-a 


I 


f  ANTERIOR.  POSTERIOR. 

I?  J  Latissimus  dorsi, ? 

O    "\  ' 

i-3    J     •    i 

(  4<  <  Pectoralis  major, Pectinaeus, 

f  Teres  major, Scansorius, 

^  j  £,  f  Subscapularis, Iliacus, 

g  J  Teres  minor, Glutaeus  minimus, 

§  |  Infra-spinatus, Glutseus  medius, 

<  [  Coraoo-brachialis, Adductors, 

Deltoid  (spinal), : Glutaeus  maximus  (sacral), 

Deltoid  (scapular), Glutaeus  maximus  (iliac), 

•  ( 

4^  •<  Supra-spinatus, Psoas  magnus, 

f  Biceps, Biceps  (ischiatic  head), 

j  ^  f  Epitrochlien, Sartorius, 

Gracilis, 


§  | Semimembranosus, 

^1 Semitendinosus, 

I"  Brachialis  anticus, Poplitseus, 
^  f  Supinator  longus, Biceps  (femoral  head), 
|  J  Pronator  radii  teres 
S  ]  Supinator  brevis, 
•<  [Pronator  quadratus, 


Triceps,  (scapular  head),  ...........................  Triceps  (rectus  or  iliac  head), 


|  |  Triceps,  (humeral  heads), Triceps  (vasti  or  femoral  heads), 

cc   C 

*Meckel,  "  Trait^  Generale  d'Anatomie  comparee."    Tome  vi.  page  413. 


w 


o 

I 


IN  THE   LIMBS  OF  MAMMALIA.  33 

ANTERIOR.  POSTERIOR. 

g>  (  Flexor  carpi  radialis  longior, Peronaeus  brevis, 

3  j,  "       brevior, Peronaeus  tertius, 

|  -j  Flexor  carpi  ulnaris, Tibialis  anticus, 

•  ^  T» 

^  (  Extensor  carpi  radialis, Peronaeus  longus, 

|  •]  «5  f  Palmaris  longus, Plantaris, 

a  <•  ^  ( "*  1 Gastrocnemius, 

2  j  ^  (  Extensor  carpi  ulnaris, Tibialis  posticus, 

O    J  f 

^Oj  \ • Soleus, 

bb  I 

§  j  Flexor  profundus  digitorum, Flexor  brevis  digitorum, 

Flexor  sublimis  digitorum, Flexor  longus  digitorum, 

4?  •]  Palmaris  brevis, 

Extensor  proprius  pollicis, Extensor  longus  digitorum, 

"         "  indicis, «          «          « 

~  ,  minimi  digiti Extensor  proprius  pollicis, 

.  /• 

Extensor  longus  digitorum, Extensor  brevis  digitorum. 

We  have  now  seen  the  general  plan,  or  the  morphology,  according  to  which  the  muscles 
of  the  mammalian  limbs  appear  to  be  arranged,  and  also  the  general  use,  or  the  physiology 
of  that  plan ;  but  in  no  case  have  we  found  it  so  closely  adhered  to  as  to  exhibit  what 
might  be  called  a  typical  condition  of  the  parts ;  everywhere  we  detect  some  variation 
according  to  the  special  functions  of  the  muscles.  The  attachments  of  a  muscle  may  be 
changed,  as  with  the  short  flexors  of  the  fingers;  but  this  is  far  more  common  with  respect 
to  the  origins  than  the  insertions,  if  indeed  the  latter  ever  are  changed ;  the  origin  of  a  mus- 
cle may  be  even  duplicated,  as  in  the  biceps  humeri.  Quite  often  there  are  extra  (acces- 
sory) muscles,  as  about  the  shoulder  arid  hip;  and  of  some  of  the  latter,  (olturatores  interims 
and  extern/us,  the  gemelli  and  quadratics  lumborum]  I  have  not  yet  been  able  to  discover  the 
morphology ;  occasionally,  too,  a  muscle  is  deficient,  at  any  rate  in  ordinary  animals,  as 
the  longitype  of  the  long  flexor  of  the  arm,  latissimus  dorsi. 

We  may  not  in  every  case  be  able  to  see  the  precise  value  of  these  variations  from  the 
general  plan ;  nor  can  we  even  generalize  them  by  asserting  that  they  are  more  numerous 
and  difficult  to  understand,  at  the  two  extremities  of  the  limbs,  as  might  be  expected  from 
the  great  mobility  allowed  by  the  ball-and-socket  joints  at  the  one,  and  the  increased  num- 
ber and  combined  movements  of  the  parts  composing  the  other,  and  that  the  muscles  are 
most  regular  in  the  intermediate  regions  of  the  limbs. 

But  whether  we  comprehend  them  or  not,  all  these  variations  must  have  a  foundation  in 
use  ;  for  all  morphology  is  for  the  sake  of  teleology ;  it  is  true  the  relation  is  also  that  of 
cause  and  effect ;  but  each  effect  becomes  in  turn  the  cause  of  some  use  below  it,  while 
each  cause  is  the  effect  of  some  power  above  it,  till  we  reach  Deity,  from  whom  all  things 
are.  In  their  highest  terms  morphology  and  teleology  are  as  Creator  and  Creation,  as  God 
and  the  Universe  :  the  one  can  only  be  manifested  through  the  other,  which  again  is  heter- 
ogeneous and  scattered  without  the  former.  It  is  the  universal  principle  of  «  concentrated 
representation ; "  many  particulars  uniting  under  one  general,  which  in  turn  unites  with 
others  under  a  higher,  and  so  on  to  Infinity. 

Morphology  and  teleology  appear  as  master  and  servant ;  but  this  is  only  an  appearance 
of  the  relation  when  viewed  a  posteriori;  and  a  view  a  priori  shows  that  the  reverse  exists 
9 


34  WILDER  ON  MORPHOLOGY  AND  TELEOLOGY 

quite  as  truly.  The  advantages  are  mutually  conferred,  and  it  is  only  a  division  of  labor. 
The  commander  of  a  company  holds  his  position  both  from  and  for  his  men,  and  only  thus 
can  several  companies  be  represented  in  a  regiment,  several  regiments  in  a  brigade,  and 
so  on  up  to  the  commander-in-chief,  who  really  represents  all  beneath  him ;  but  what  is 
he  without  them  ?  Every  superior  is,  or  ought  to  be,  more  truly  the  servant  of  his  infe- 
riors, who  while  they  appear  to  obey  him  really  serve  themselves.  Broad  as  is  our  land,  it 
cannot  limit  the  application  of  our  national  motto.  "  E pluribus  unum"  is  not  merely  a 
national  motto,  but  the  concise  expression  of  an  all-pervading  law,  the  basis  of  the  highest 
natural,  human,  and  Divine  order. 

It  may  now  be  asked,  Which  is  of  the  greater  importance,  and  deserving  of  the  more  at- 
tention,— morphology  or  teleology,  the  general  laws,  or  the  particular  facts  and  uses  which 
they  represent  ?  The  answer  to  this  question  will  vary  with  the  three  degrees  of  depth  to 
which  our  search  is  carried.  At  first  we  exclaim,  "  The  facts,  of  course ;  they  only  have 
an  actual  existence,  and  are  of  any  real  use ;  they  must  be  diligently  collected  and  examined  ; 
they  are  strange  and  beautiful,  and  our  wonder  and  admiration  are  constantly  aroused." 
But  there  is  something  beyond  this.  The  facts  resemble  each  other,  some  more,  some 
less,  and  soon  we  arrange  them  in  groups,  acknowledging,  if  such  grace  be  given  us,  that 
those  groups  really  did  exist  before  we  saw  them ;  our  minds  are  occupied  with  these ; 
we  give  them  names,  and  delight  in  contemplating  the  laws  and  principles  they  suggest 
to  us.  We  find,  moreover,  that,  though  immaterial,  they  have  a  most  substantial  mental  exist- 
ence, and  now  we  accord  to  this  study  the  higher  importance,  and,  mounted  upon  our 
philosophical  superstructure,  are  inclined  utterly  to  ignore  the  groundwork. 

It  is  as  if  one  had  labored  long  in  piling  stones  together  to  build  a  lofty  tower,  and  at  last 
standing  upon  the  single  block  which  forms  the  summit,  forgot  all  below,  acknowledging 
only  himself  and  the  result  of  his  own  work.  If  he  stays  there,  it  is  clear  that  he  can  be 
of  little  use  to  himself  or  to  any  one  else  ;  he  must  descend  and  show  to  others  the  way  up. 
And  now  if  he  does  this, — if  he  has  employed  his  temporary  elevation  in  looking  abroad  and 
beneath  as  well  as  above,  and,  more  than  all,  if  he  imparts  to  others  the  superior  informa- 
tion thus  acquired,  and  instructs  them  that  they  also  may  ascend,  then  he  has  accomplished 
far  more  than  if  he  had  remained  below  surrounded  and  overwhelmed  by  the  unarranged, 
and  therefore  uninstructive  abundance,  or  had  stayed  at  the  top,  proud  and  disdainful  of 
those  beneath  him;  and  he  now  perceives  that  the  former  position  was  undesirable  and  the 
latter  impossible  without  the  other,  and  accords  to  each  its  true  value  in  what  h£  can  accom- 
plish with  their  combined  assistance. 

These  three  states  of  mind  are  respectively  those  of  the  unthinking  but  observant  child, 
of  the  reasoning  philosophical  youth,  and  of  the  wise  man  who,  having  passed  through 
both  these  stages,  has  attained  to  something  better  than  either, — a  power  and  a  disposition 
to  use  what  he  has  gained  for  others. 

Three  states  are  mentioned.  There  is  really  a  fourth,  but  it  is  the  first  in  the  series,  and 
corresponds  to  the  embryo,  which  manifests  no  life,  and  is  as  it  were  the  ground  in  which 
are  implanted  the  others  in  their  order.  It  is  the  stage  of  inactivity,  of  preparation,  and 
it  is  easy  to  see  the  analogy  between  this  and  the  lowest  sub-kingdom  of  animals.  The 
first  and  the  last  states  seem  in  a  measure  to  resemble  each  other  on  a  lower  and  higher 
plane,  as  the  vertebrate  type  stands  over  the  radiate.  And  as  the  mature  animal  and  the 
full-grown  tree,  in  all  their  strength  and  beauty,  expend  their  best  energies  in  the  elabora- 
tion of  just  such  simple  eggs  and  seeds  as  those  from  which  they  sprung,  so  the  latter  are 
the  morphological  epitomes  of  what  may  be,  the  other  teleological  expressions  of  what  has 
been. 


IN  THE  LIMBS  OF  MAMMALIA.  05 

Among  students  of  Nature,  the  three  latter  states  of  mind  are  respectively  those  of  the 
mere  collector  or  dissector ;  of  the  votary  of  morphology  and  classification  alone ;  and 
lastly,  of  the  favored  few  who  happily  combine  them  both,  and  thus  accomplish  more  than 
with  either  one  alone ;  and  can  we  not  see  that  the  industry  which  has  succeeded  in  accu- 
mulating such  vast  numbers  of  facts  is  already  giving  way  to  philosophical  reasonings  and" 
a  clearer  comprehension  of  the  same?  The  last  stage  of  science  is  one  to  be  striven  for 
with  full  belief  in  its  existence  in  the  future. 

ADDENDUM.     For  extended  definition  and  illustration  of  Teleology,  see  the  chapter  on  Teleology  of  the  Skele- 
ton of  Fishes,  in  Owen's  Comparative  Anatomy,  Vol.  ii. 

NOTE.     The  foregoing  paper,  in  a  much  less  extended  form,  but  containing  most  of  the  principal  ideas,  was  ori<n- 
nally  prepared  and  presented  as  a  Thesis  at  my  graduation  in  the  Department  of  Comparative  Anatomy  and  Physi 
ology  in  the  Lawrence  Scientific  School  of  Harvard  University  in  July,  1862.     Since  its  revision  I  have  received 
from  Professor  James  D.  Dana  copies  of  the  following  papers  by  him : 

1.  «  On  Cephalization."     From  the  "  New  Englander  "  for  July,  1863. 

2.  «  On  Parallel  Relations  of  the  Classes  of  Vertebrates,  and  on  some  Characteristics  of  the  Reptilian  Birds  " 
From  the  American  Journal  of  Science  and  Arts,  Vol.  xxxvi.  Nov.  1863. 

3.  "The  Classification  of  Animals,  based  on  the  Principles  of  Cephalization."     Am.  Journ.  Science  etc    Vol. 
xxxvi.  Nov.  1863,  and  xxxvii.  Jan.  1864. 

4.  «  On  Fossil  Insects  from  the  Carboniferous  Formation  in  Illinois."    Am.  Journ.  Science,  etc.,  Vol.  xxxvii 
Jan.  1864. 

Also,  from  Norton  Folsom,  M.  D.,  Surgeon  of  the  45th  Regiment  U.  S.  colored  troops,  the  manuscript  notes  of 
,ssay  by  him  «on  Anatomical  Symmetry,"  read  at  the  Commencement  of  the  Massachusetts  Medical  College  in 
July,  1863. 

All  these  papers  I  have  read  with  great  interest  and  pleasure,  not  only  for  their  intrinsic  scientific  value  but  also 
;ause  in  some  portions  of  them  were  contained  confirmations  of  the  ideas  expressed  in  this  paper,  which  confir- 
mations are  the  more  gratifying  as  coming  from  such  masters  in  the  science  as  Wyman  and  Dana,  from  the  former 
of  whom,  Dr.  Folsom  writes,  many  of  the  ideas  in  his  essay  were  derived. 

I  write  this  in  order  that  the  coincidences  between  the  views  in  the  papers  above  mentioned  and  my  own  may 
5  lessen  the  originality  of  what  was  written  some  months  before  those  papers  were  read  by  me. 

BURT  G.  WILDER, 

Surgeon  55th  Mass.  Vol  Infantry. 
CHARLESTON,  S.  C.,  August  llth,  1865. 

Published,  November,  1865. 


14  DAY  USE 

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